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Bulletin  of  the 

British  Museum  (Natural  History) 


^36 

PRESENTED 

' 


A  review  of  the  Miletini 
(Lepidoptera:  Lycaenidae) 

J.  N.  Eliot 


Entomology  series 
Vol53  Nol 


25  September  1986 


The  Bulletin  of  the  British  Museum  (Natural  History),  instituted  in  1949,  is  issued  in  four 
scientific  series,  Botany,  Entomology,  Geology  (incorporating  Mineralogy)  and  Zoology, 
and  an  Historical  series. 

Papers  in  the  Bulletin  are  primarily  the  results  of  research  carried  out  on  the  unique  and 
ever-growing  collections  of  the  Museum,  both  by  the  scientific  staff  of  the  Museum  and  by 
specialists  from  elsewhere  who  make  use  of  the  Museum's  resources.  Many  of  the  papers  are 
works  of  reference  that  will  remain  indispensable  for  years  to  come. 

Parts  are  published  at  irregular  intervals  as  they  become  ready,  each  is  complete  in  itself, 
available  separately,  and  individually  priced.  Volumes  contain  about  300  pages  and  several 
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Publications  Sales, 

British  Museum  (Natural  History), 
Cromwell  Road, 

London  SW75BD, 
England. 


World  List  abbreviation:  Bull.  Br.  Mus.  nat.  Hist.  (Ent.) 


Tustees  of  the  British  Museum  (Natural  History),  1986 


The  Entomology  series  is  produced  under  the  general  editorship  of  the 

Keeper  of  Entomology:  Laurence  A.  Mound 

Assistant  Editor:  W.  Gerald  Tremewan 


ISBN  0  565  06019  8 
ISSN  0524-6431 


British  Museum  (Natural  History) 
Cromwell  Road 
London  SW7  5BD 


Entomology  series 
Vol53Nolppl-105 


Issued  25  September  1986 


(NATURAL  HISTORY) 

19 

PRESENTED 
GENr 


Bulletin  of  the 

British  Museum  (Natural  History) 


Entomology  series    Vol  53     1986 


British  Museum  (Natural  History) 
London  1986 


Dates  of  publication  of  the  parts 

No  1                 25  September  1986 

No  2        .        .        .        . 30  October  1986 

No  3        .        .        .        .        .        .        .        .        .        .        .  30  October  1986 

No  4 27  November  1986 

No  5  18  December  1986 


ISSN  0524-6431 


Printed  in  Great  Britain  by  Henry  Ling  Ltd.,  at  the  Dorset  Press,  Dorchester,  Dorset 


Contents 
Entomology  Volume  53 


No  1       A  review  of  the  Miletini  (Lepidoptera:  Lycaenidae). 

J.N.Eliot 1 

No  2       Australian  ichneumonids  of  the  tribes  Labenini  and  Poecilocryptini. 

I.  D.  Gauld  &  G.  A.  Holloway ...       107 

No  3       The  tribe  Pseudophloeini  (Hemiptera:  Coreidae)  in  the  Old  World  tropics  with 
a  discussion  on  the  distribution  of  the  Pseudophloeinae. 
W.  R.  Dolling 151 

No  4      The  songs  of  the  western  European  grasshoppers  of  the  genus  Omocestus  in 
relation  to  their  taxonomy  (Orthoptera:  Acrididae). 
D.  R.  Ragge 213 

No  5      The  structure  and  affinities  of  the  Hedyloidea:  a  new  concept  of  the  butterflies. 

M.J.  Scoble.  251 


A  review  of  the  Miletini  (Lepidoptera:  Lycaenidae) 

J.  N.  Eliot 

Upcott  House,  Bishop's  Hull,  Taunton,  Somerset  TA4  1AQ 

Contents 

Synopsis 1 

Introduction 1 

Acknowledgements 2 

Checklist  of  the  Miletini 3 

Tribal  characters 6 

Key  to  the  genera  of  Miletini 7 

Genus  AllotinusC.&R.Felder 7 

Genus  Logania  Distant 57 

Genus  Lontalius  gen.  n 74 

Genus  Miletus  Hiibner 75 

Genus  Megalopalpus  Rober 84 

References 86 

Index 104 

Synopsis 

This  review  of  the  Miletini  is  based  mainly  on  characters  of  the  male  genitalia.  Keys  to,  and  descriptions  of, 
the  genera,  subgenera,  species  and  subspecies  are  provided.  One  genus,  one  subgenus,  eight  species  and 
seven  subspecies  are  newly  described.  One  species  is  relegated  to  subspecies,  and  three  subspecies  are 
raised  to  species;  41  new  synonyms  are  established. 

Introduction 

The  butterflies  which  are  here  considered  to  comprise  the  Oriental  section  of  the  tribe  Miletini 
Reuter,  1897  (see  Corbet,  1939b)  were  originally  given  family-group  status  (as  the  Gerydinae) 
by  Doherty  (1886),  mainly  on  the  basis  of  their  distinctive  eggs.  Reuter  (1897:  263)  included  the 
African  genus  Megalopalpus  Rober,  1886,  in  the  group,  treating  the  latter  as  a  tribe,  the 
Miletidi,  which  is  distinguished  from  all  other  Lycaenidae  by  the  peculiar  labial  palpi  bearing 
long  hairs  on  the  inner  surface  of  the  basal  segment  (Basalfleck).  Corbet  (19396)  used  the  family 
group  name  Miletinae  in  place  of  Gerydinae  because  Gerydus  Boisduval,  1836,  is  an  objective 
synonym  of  Miletus  Hiibner,  1819;  his  action  is  valid  under  Article  40b  of  the  International  Code 
of  Zoological  Nomenclature  (1985).  Eliot  (1973)  used  Miletinae  in  a  wider  sense  than  previous 
authors,  placing  the  Miletini  (Miletinae  sensu  Corbet)  as  one  of  four  tribes  which  he  included  in 
the  subfamily. 

Only  two  complete  analyses  of  the  Oriental  Miletini  (sensu  Eliot)  have  so  far  been  attempted, 
both  by  Fruhstorfer  who  based  his  studies  in  part  on  a  number  of  genitalia  preparations  made  by 
Reverdin.  In  the  first  (Fruhstorfer,  1913-14)  it  is  evident  that  the  new  taxa  he  described  had 
three  separate  origins.  Many  descriptions  are  based  on  specimens  in  his  own  collection,  some  of 
which  bear  type  and/or  determination  labels  attached  either  at  that  time  or  at  some  later  date. 
Others  relate  to  specimens  in  the  collections  of  the  British  Museum  (Natural  History)  and 
Walter  Rothschild  at  Tring,  to  which  no  type  or  determination  labels  were  attached;  these 
collections  were  studied  by  Fruhstorfer  during  a  visit  to  England.  Finally,  a  few  descriptions  are 
based  on  records  and  figures  by  other  authors  of  specimens  which  he  had  not  seen;  in 
Fruhstorfer's  first  analysis  such  taxa  were,  in  most  cases,  based  on  a  work  by  Semper  (1889),  so 
those  specimens  listed  by  Semper  may  be  considered  to  comprise  Fruhstorfer's  type-series.  In 
certain  cases,  however,  it  is  clear  that  Fruhstorfer  applied  a  name  to  a  single  specimen  figured  by 


Bull.  Br.  Mus.  not.  Hist.  (Ent.)  53  (1):  1-105  Issued  25  September  1986 


2  J.  N.   ELIOT 

Semper,  and  in  such  cases  the  original  of  the  figure  is  automatically  the  holotype;  in  other  cases, 
however,  it  has  been  necessary  to  designate  a  lectotype.  In  his  second  analysis  (Fruhstorfer, 
1916)  it  is  evident  that  he  had  in  the  meantime  examined  Semper's  collection,  and  he  described 
further  material  to  which  type  and  determination  labels  were  attached.  There  is  some  evidence 
to  suggest  that  during  this  examination  some  of  Semper's  original  labels  were  inadvertently 
transferred  from  one  specimen  to  another.  Such  instances  are  mentioned  later  under  the 
relevant  taxon. 

As  Fruhstorfer  seldom  selected  types,  I  have  designated  lectotypes  when  it  is  not  clear  that  a 
taxon  was  described  from  a  single  specimen.  I  have  also  designated  lectotypes  for  some  taxa 
named  by  other  authors  when  I  have  been  able  to  recognise  primary  type-material  in  the  British 
Museum  (Natural  History).  When  the  original  type-series  are  in  museums  which  I  have  not 
visited,  I  have  generally  left  the  selection  of  lectotypes  to  future  workers,  except  in  a  few  cases 
where,  to  avoid  confusion,  I  have  designated  figured  specimens. 

The  following  abbreviations  have  been  used  for  the  actual  or  supposed  location  of  type- 
material. 

BMNH  British  Museum  (Natural  History),  London 

CM  Carnegie  Museum,  Pittsburgh 

MNHU  Museum  fiir  Naturkunde  der  Humboldt-Universitat,  Berlin 

RNH  Rijksmuseum  van  Natuurlijke  Historic,  Leiden 

SM  Senckenberg  Museum,  Frankfurt  am  Main 

SMT  Staatliches  Museum  fiir  Tierkunde,  Dresden 

ZSI  Zoological  Survey  of  India,  Calcutta 

Here  it  may  be  noted  that  there  are  in  the  BMNH  collection  a  number  of  specimens  bearing 
red  type  labels  as  well  as,  in  some  cases,  labels  in  Corbet's  handwriting  identifying  them  as 
holotypes  or  allotypes  which  were  not  cited  as  such  in  the  original  descriptions.  Where  it  is 
reasonable  to  assume  that  these  were  part  of  the  original  type-series  I  have  normally  designated 
them  as  lectotypes  in  preference  to  any  other  syntypes.  In  some  cases,  however,  it  is  quite  clear 
that  the  specimens  cannot  possibly  be  part  of  the  original  type-series;  such  specimens  I  have 
rejected  as  syntypic  material  and  labelled  them  accordingly.  These  'types'  are  listed,  however,  in 
the  section  dealing  with  species  and  subspecies  since  it  is  possible  that  they  may  have  misled 
authors  in  the  past  or  might  do  so  in  the  future. 

Acknowledgements 

This  revision  is  based  mainly  on  material  in  the  BMNH,  and  I  am  grateful  to  the  Trustees  for  access  to  the 
collections  and  library,  to  Mr  R.  I.  Vane-Wright  and  Mr  P.  R.  Ackery  for  their  help  and  encouragement, 
and  to  the  Photographic  Unit  for  providing  the  photographs  for  Figs  55-104.  I  thank  especially  Mr 
Vane- Wright  and  Mrs  S.  M.  North  for  undertaking  a  review  of  the  African  genus  Megalopalpus,  of  which  I 
have  no  practical  knowledge  in  the  field,  and  for  preparing  the  draft  incorporated  in  this  paper.  I  am  also 
very  grateful  to  Dr  Heinz  Schroder  for  the  loan  of  important  material  from  the  Semper  collection  in  the 
Senckenberg  Museum,  without  which  it  would  have  been  impossible  to  carry  out  a  satisfactory  review.  I 
thank  Dr  R.  de  Jong,  RNH,  and  Professor  S.  Murayama,  who  sent  material  for  examination.  Mr  G.  C. 
Treadaway  placed  the  whole  of  his  extensive  Philippine  material  at  my  disposal  and  sent  me  the 
photographs  for  Figs  105-108.  I  also  received  valuable  Philippine  material  through  the  generosity  of  Mr 
Yusuke  Takanami  and  Dr  A.  Ballantine,  including  new  taxa  which  I  have  named  in  their  honour.  Major  A. 
Bedford  Russell  lent  me  material  from  Vietnam  and  Sulawesi,  and  presented  the  holotype  of  Allotinus 
samarensis  russelli  to  the  BMNH,  and  Dr  E.  Diehl  sent  me  material  from  Simeulue.  In  addition  the 
collections  of  the  late  Mr  W.  A.  Fleming  (kindly  placed  at  my  disposal  by  his  widow),  Lt-Col  C.  F.  Cowan, 
Dr  T.  Norman  and  my  own  have  been  examined. 


THE  MILETINI 


Checklist  of  the  Miletini 


Genus  ALLOTINUS  C.  &  R.  Felder,  [1865] 

Subgenus  ALLOTINUS  C.  &  R.  Felder,  [1865] 
fallax  C.&R.  Felder,  [1865] 
fallaxfallaxC.  &  R.  Felder,  [1865] 
sabazus  Fruhstorfer,  1913  syn.  n. 
zaradrus  Fruhstorfer,  1916 
fallax  aphacus  Fruhstorfer  ,1913 
ancius  Fruhstorfer,  1913  syn.  n. 
artinus  Fruhstorfer,  1916 
fallax  eryximachus  Fruhstorfer,  1913 
fallax  dotion  Fruhstorfer,  1913 
fallax  tymphrest us  Fruhstorfer,  1916 
fallax audaxH.  H.  Druce,  1895 
fallax  apusde  Niceville,  1895 

michaelis  Eliot,  1959 
albifasciatus  Eliot,  1980 
subvioIaceusC.  &  R.  Felder,  [1865] 
subvioIaceussubviolaceusC.  &  R.  Felder, 
[1865] 

alkamah  Distant,  1886  syn.  n. 
manychus  Fruhstorfer,  1913  syn.  n. 
kallikrates  Fruhstorfer,  1913  syn.  n. 
silarus  Fruhstorfer,  1916  syn.  n. 
subviolaceusmirusVan  Eecke,  1914 
agnolia  sp.  n. 
nicholsi  Moulton  ,1911 

nicholsi  nicholsi  Moulton,  191 1 
nicholsi  battakanus  Fruhstorfer,  1913 
major  C.  &  R.  Felder,  [1865] 

depictus  Fruhstorfer,  1913  syn.  n. 
kdldwarus  Ribbe,  1926 
max/musStaudinger,  1888  stat.  n. 

Subgenus  FABITARAS  subgen.  n. 
fabius  (Distant  &  Pryer,  1887) 
fabiusfabius  (Distant  &  Pryer,  1887) 
caudatus  Grose-Smith,  1893  syn.  n. 
pamisus  Fruhstorfer,  1914  syn.  n. 
fa  biusarrius  Fruhstorfer,  1914 
borneensis  Moulton,  1911 

elioti  Corbet,  1939  syn.  n. 
punctatus  (Semper,  1889) 

anaxandridas  Fruhstorfer,  1916  syn.  n. 
caesemius  Fruhstorfer,  1916  syn.  n. 
nigritus  (Semper,  1889) 

eretria  Fruhstorfer,  1916 
strigatus  Moulton,  1911 

strigatus  strigatus  Mouhon,  1911 
strigatus  malayanus  Corbet ,  1939 
denalus  Corbet,  1939  syn.  n. 
brooks!  sp.  n. 
bidiensissp.  n. 
faras(Doherty,  1889) 

panormis  (Elwes,  1893)  syn.  n. 
sarrastes  Fruhstorfer,  1913  stat.  n. 
mendava  Riley,  1944  syn.  n. 
porriginosus  Toxopeus,  1932  syn.  n. 


portunus  (de  Niceville,  1894) 

portunus port unus  (de  Niceville,  1894) 

narsares  Fruhstorfer,  1913  syn.  n. 
portunus  maitus  Fruhstorfer,  1914 

fruhstorferi  Corbet,  1939  syn.  n. 
portunus pyxus  (de  Niceville,  1894) 

wdterstradti  H.  H.  Druce,  1895 

Subgenus  PARAGERYDUS  Distant,  1884 
Miletographa  Rober,  1892 
horsffeldi  (Moore,  1857) 
horsfieldi  horsfieldi  (Moore ,  1857) 
horsfieldi  permagnus  Fruhstorfer,  1913 
infumata  Fruhstorfer,  1913 
nessus  Corbet,  1939 
horsfieldi  siporanus  Riley,  1944 
horsfieldi  satelliticus  Fruhstorfer,  1913 
leogoron  Fruhstorfer,  1916 

leogoron  leogoron  Fruhstorfer,  1916 

intricata  Fruhstorfer,  1913  (unavailable  name) 
vadosus  Corbet,  1939 
lindus  Corbet,  1939 
leogoron  normani  Eliot,  1967 
leogoron  batuensis  Eliot,  1967 
leogoron  plessis  Eliot,  1967 
melos  (H.  H.  Druce,  1896)  sp.  rev. 
reverdini  Fruhstorfer,  1916  syn.  n. 
talu  Eliot,  1967  syn.  n. 
samarensissp.  n. 
samarensis  samarensis  subsp.  n. 
samarensis  russelli  subsp.  n. 
macassarensis  (Holland,  1891) 
macassarensis  macassarensis  (Holland,  1891) 

damodar  Fruhstorfer,  1913 
macassarensis  menadensis  Eliot,  1967 
luzonensis  Eliot,  1967  stat.  n. 
albatusC.  &  R.  Felder,  [1865] 

albatusalbatusC.  &  R.  Felder,  [1865] 
;i//>;ifi/,v/iiriH/;/vsiihsp.  n. 
apries  Fruhstorfer,  1913 

apriesapries  Fruhstorfer,  1913 

eupalion  Fruhstorfer,  1914  syn.  n. 
apries  dositheus  Fruhstorfer,  1914 
apries  ristus  subsp.  n. 
corbeti  Eliot,  1956 
unicolorC.  &  R.  Felder,  [1865] 

unicolorunicolorC.  &  R.  Felder,  [1865] 
eurytanus  Fruhstorfer,  1913  syn.  n. 
dilutus  Corbet,  1939  syn.  n. 
unicolor  continental  Fruhstorfer,  1913 

dtddnus  Fruhstorfer,  1916 
unicolor  rekkia  Riley  &  Godfrey,  1920 
unicolor mooreiH.  H.  Druce,  1895 

rebilus  Fruhstorfer,  1913  syn.  n. 
unicolor  aphocAaKheil,  1884 

myridndus  Fruhstorfer,  1913  syn.  n. 
unicolor  posidion  Fruhstorfer,  1913 

molionides  Fruhstorfer,  1913  syn.  n. 


J.  N.  ELIOT 


niceratus  Fruhstorfer,  1913  syn.  n. 
bajanus  Fruhstorfer,  1913  syn.  n. 
enatheus  Fruhstorfer,  1913 
suka  Piepers  &  Snellen,  1918 
enganicus  Fruhstorfer,  1913  syn.  n. 
unicolor  georgius  Fruhstorfer,  1913 

leitus  Fruhstorfer,  1916 
unicolor  zitema  Fruhstorfer,  1916 
paetus(de  Niceville,  1895) 
parapus  Fruhstorfer,  1913 
nivalis  (H.  Druce,  1873) 
nivalis  nivalis  (H.  Druce,  1873) 
nivalis  felderi  Semper ,  1 889 
substrigosus  (Moore,  1884) 
substrigosus  substrigosus  (Moore,  1884) 

magaris  Fruhstorfer,  1913 
substrigosus  lenaia  Fruhstorfer,  1913 
substrigosus  sibyllinus  Riley,  1944 
substrigosus  ballantinei  subsp.  n. 
substrigosus  yusukei  subsp.  n. 
da  vidis  Eliot,  1959 
(I  mini  In  (Moore,  [1866]) 
drumila  drumila  (Moore,  [1866]) 
multistrigatus  de  Niceville,  1886 
insignis  (Staudinger,  1888) 
drumila  aphthonius  Fruhstorfer,  1913 
grisea  Riley  &  Godfrey,  1920 

Genus  LOGANIA  Distant,  1884 
Malais  Doherty,  1889 
malayica  Distant,  1884 
malayica  malayica  Distant,  1884 
malayica  subura  Fruhstorfer,  1914 
nehalemia  Fruhstorfer,  1914  stat.  rev. 
waltraudae  sp.  n. 
regina  (H.  Druce,  1873) 
regina  regina  (H.  Druce,  1873) 

evora  Fruhstorfer,  1916  syn.  n. 
regina  sriwa  Distant,  1886 
paluana  sp.  n. 
marmorata  Moore,  1884 
marmorata  marmorata  Moore,  1884 
marmorata  damis  Fruhstorfer,  1914 
marmorata  hilaeira  Fruhstorfer,  1914 
obscura  Distant  &  Pryer,  1887 

(nom.  preocc.) 
nada  Fruhstorfer,  1914 
stenosa  Fruhstorfer,  1916  syn.  n. 
cineraria  Fruhstorfer,  1916  syn.  n. 
sora  Fruhstorfer,  1916  syn.  n. 
marmorata  lahomius(K.hei\,  1884) 
marmorata  diehJi  subsp.  n. 
marmorata  munichya  Fruhstorfer,  1914 
marmorata  javanica  Fruhstorfer,  1914 

glypha  Fruhstorfer,  1914 
marmorata  palawana  Fruhstorfer,  1914 

distanti  Staudinger,  1889  (nom.  preocc.) 
marmorata  samosata  Fruhstorfer,  1914 
marmorata  faustina  Fruhstorfer,  1914 


obscura  Rober,  1886 

martinus  (Fruhstorfer,  1913)  syn.  n. 
donussa  Fruhstorfer,  1916  syn.  n. 
distanti  Semper,  1889 

distanti  distanti  Semper ,  1889 
apsines  Fruhstorfer,  1914 
turdeta  Fruhstorfer,  1916  syn.  n. 
distanti massalia  Doherty,  1891  stat.  n. 

luca  de  Niceville,  1894  syn.  n. 
distanti  drucei  Moulton  ,1911 
distanti  staudingeriH.  H.  Druce,  1895 
hampsoni  Fruhstorfer,  1914 

meeki  Rothschild,  1915  syn.  n. 
masana  Fruhstorfer,  1916  syn.  n. 
watsoniana  de  Niceville,  1898 
subfasciata  Tytler,  1915 

Genus  LONTALIUS  gen.  n. 
eltussp.  n. 
eltus  eltus  subsp.  n. 
eltus  treadawayi  subsp.  n. 

Genus  MILETUS  Hubner,  1819 
Symetha  Horsfield,  1828 
Gerydus  Boisduval,  1836 
Archaeogerydus  Fruhstorfer,  1916 
chinensisC.  Felder,  1862 

chinensischinensisC.  Felder,  1862 

chinensislearchusC.  &  R.  Felder,  [1865] 
irroratusH.  Druce,  1874 
kelantanus  Corbet,  1938 

chinensis  assamensis  (Doherty,  1891) 
milvius  (Fruhstorfer,  1913) 

chinensis  longeana  (de  Niceville,  1898) 
croton  (Doherty,  1889) 

croton  croton  (Doherty,  1889) 
tavoyana  (Evans,  1932) 

croton  corus  Eliot,  1961 

croton  karennius  (Evans,  1932) 
mallus  (Fruhstorfer,  1913) 

mallusmallus  (Fruhstorfer,  1913) 

mallus  gethusus  (Fruhstorfer,  1913) 

mallus  shanius  (Evans,  1932) 
gaesa  (de  Niceville,  1895) 

gaesagaesa  (de  Niceville,  1895) 

gaesa  carrinas  (Fruhstorfer,  1916) 
nymphis  (Fruhstorfer,  1913) 

n ymphis  n ymphis  (Fruhstorfer,  1913) 

nymphis  porus  Eliot ,  1 96 1 

nymphis  fictus  Corbet,  1939 

nymphis  eneus  Eliot,  1961 
zinckeniiC.  &  R.  Felder,  [1865] 

zinckeniizinckeniiC.  &  R.  Felder,  [1865] 

zinckenii improbus  (H.  H.  Druce,  1895) 
gopara  (de  Niceville,  1890) 

goparagopara  (de  Niceville,  1890) 
denticulata  (Fruhstorfer,  1913) 

gopara  pardus  Eliot ,  196 1 

gopara  eustatius  (Fruhstorfer,  1913) 


THE  MILETINI 


gopara  artaxatus  (Fruhstorfer,  1913) 

oichalia  (Fruhstorfer,  1913) 
valeus  (Fruhstorfer,  1913) 

pallaxopas  (Fruhstorfer,  1913) 
gaetulus  (de  Niceville,  1894) 

gaet ulus gaet ulus  (de  Niceville,  1894) 

gaetulus innocens(H.  H.  Druce,  1895) 

gaetulus  aphytis  (Fruhstorfer,  1913) 
boisduvali  Moore,  1857 

boisduvali  boisduvali  Moore,  1857 
vincula  (H.  H.  Druce,  1895) 
heraeon  (Fruhstorfer,  1916) 
courvoisieri  (Fruhstorfer,  1915) 
oxylus  (Fruhstorfer,  1916) 
lombokianus  (Fruhstorfer,  1913) 
acragas  (Doherty,  1891) 
buruensis  (Holland,  1900) 
ceramensis  Ribbe,  1889 
dossemus  (Fruhstorfer,  1913) 
stygianus  Butler,  1884 
adeus  (Fruhstorfer,  1913) 

boisduvali  diotrophes  (Fruhstorfer,  1913) 

boisduvali  avitus  (Fruhstorfer,  1916) 
drucei  (Semper,  1889) 

druceidrucei  (Semper,  1889) 
philippus  (Staudinger,  1889) 
jacchus  (Fruhstorfer,  1913) 
palanius  (Fruhstorfer,  1913) 
epidurus  (Fruhstorfer,  1913) 

drucei  metrovius  (Fruhstorfer,  1913) 
phradimon  (Fruhstorfer,  1915) 
biggsii  (Distant,  1884) 

biggsii  biggsii  (Distant,  1884) 
atomaria  (Fruhstorfer,  1913) 
xeragis  (Fruhstorfer,  1916) 
hyllus  (Fruhstorfer,  1916) 
sebethus  (Fruhstorfer,  1916) 
extraneus  (Toxopeus,  1929) 

biggsii  natunensis  (Fruhstorfer,  1916) 

biggsii  niasicus  (Fruhstorfer,  1913) 
batunensis  (Fruhstorfer,  1913) 

biggsii  albotignula  (Van  Eecke,  1914) 

simalurensis  (Toxopeus,  1928) 
cellarius  (Fruhstorfer,  1913) 
symethus  (Cramer,  1779) 

symethus  symethus  (Cramer,  1779) 
pandu  (Horsfield,  1828) 

symethus  petronius  (Distant  &  Pryer,  1887) 
diopeithes  (Fruhstorfer,  1913) 
bangkanus  (Fruhstorfer,  1914) 
hieropous  (Fruhstorfer,  1916) 

symethus  solitariusOkubo,  1983 

symethus  acampsis  (Fruhstorfer,  1913) 

symethus  nuct us  Eliot,  1961 

symethus  perlucidus  (Fruhstorfer,  1913) 
megaris  (Fruhstorfer,  1913) 

symethus  vespesianus  (Fruhstorfer,  1913) 

symethus  batuensis  (Fruhstorfer,  1914) 


symethus  edonus  (Fruhstorfer,  1913) 
symethus  philopator  (Fruhstorfer,  1914) 
symethus  hierophantes  (Fruhstorfer,  1916) 
symethus  phant us  subsp.  n. 

gallus  (de  Niceville,  1894) 
gallus  gallus(de  Niceville,  1894) 
gallus  leucocyon  (Toxopeus,  1940) 
heracleion  (Doherty,  1891) 

heracleion  heracleion  (Doherty,  1891) 
heracleion  arion  Eliot,  1961 
a/icon  (Doherty,  1889) 
ancon  ancon  (Doherty,  1889) 
ancon  gigas(H.  H.  Druce,  1895) 

anconides  (Fruhstorfer,  1913) 
archilochus  (Fruhstorfer,  1913) 

archilochus  archilochus  (Fruhstorfer,  1913) 
archilochus  siamensis  (Godfrey,  1916) 
gigantes(de  Niceville,  1894) 
atimonicus  Murayama  &  Okamura,  1973 
celinus  Eliot,  1961 
takanamiisp.  n. 
/eos(Guerin-Meneville,  1830) 
leosteos  (Doherty,  1891) 
leos  fforensis  (Fruhstorfer,  1913) 

eulus  (Fruhstorfer,  1913) 
leostellus  (Fruhstorfer,  1913) 
leos  catoleucos  (Fruhstorfer,  1913) 
leos  maximus  (Holland ,  1 89 1 ) 
divisa  (Fruhstorfer,  1913) 
sarus  (Fruhstorfer,  1913) 
leosvaneecki  (Toxopeus,  1930) 
leos  mangolicus  (Fruhstorfer,  1913) 
/eos/eos(Guerin-Meneville,  1830) 
boisduvalii  (Butler,  1884) 
meronus  (Fruhstorfer,  1913) 
amphiarus  (Fruhstorfer,  1913) 
gardineri  (Fruhstorfer,  1914) 
leos  virtus  (Fruhstorfer,  1913) 

pentheus  (Fruhstorfer,  1913) 
leosaronicus  (Fruhstorfer,  1914) 
nineyanus  (Fruhstorfer,  1914) 
acrisius  (Fruhstorfer,  1914) 
melanion  C.  &  R.  Felder,  [1865] 

melanionmelanionC.  &  R.  Felder,  [1865] 

albiguttatus  f.  n.  (infrasubspecific  name) 
melanion  euphranor  (Fruhstorfer,  1914) 
bazilan us  (Fruhstorfer,  1913)  stat.  n. 

vitelianus  (Fruhstorfer,  1913)  syn.  n. 

Genus  MEGALOPALPUSRober,  1886 

angulosus  Griinberg,  1910 
mete/eucusKarsch,  1893 
simplex  Rober,  1886 

bicoloria  (Capronnier,  1889) 

similis  (Kirby,  1890) 

gigas  Bethune-Baker,  1914 
zymna  (Westwood,  1851) 

pallida  Aurivillius,  1922 


6  J.   N.  ELIOT 

Tribal  characters 

The  characters  of  Miletini  (sensu  Eliot,  1973)  are  as  follows.  Eyes  smooth.  Antennae  with 
narrow,  gradually  incrassate  club  and  with  the  nudum  extending  down  the  shaft  to  the  base  or 
very  nearly  so.  Labial  palpi  asymmetrical,  but  there  is  no  constancy  in  which  palpus  is  the  longer. 
In  most  species  the  palpi  are  unusually  long  and  thin,  more  so  in  females  than  in  males,  and 
protrude  well  beyond  the  head;  they  are  also  unusual  in  that  the  'Basalfleck'  of  Reuter  is  clothed 
with  hairs  of  unknown  function.  Proboscis  long,  bearing  many  sensilla  throughout  its  length. 
Legs  more  or  less  abnormal,  with  the  tarsi  very  long,  flattened  and  blade-like  in  Miletus  and 
Megalopalpus,  cylindrical  but  very  long  and  thin  in  Allotinus  and  with  the  tibiae  outwardly 
swollen  or  incrassate  in  Logania  and  Lontalius.  The  mid-  and  hind-tibiae  lack  the  usual  pair  of 
terminal  spurs.  The  male  fore-tarsus  is  reduced  to  a  single  segment  ending  abruptly  but  with  a 
small  pointed  process  directed  downwards,  except  in  the  typical  species  of  Logania  in  which  the 
tarsus  tapers  to  a  down-curved  point.  The  claws  on  the  mid-  and  hind-tarsi  of  both  sexes  are 
small,  and  minute  on  the  fore-tarsi  of  females.  The  abdomens  of  males  are  long  and  protrude 
well  beyond  the  hindwings,  except  in  Allotinus  major.  There  is  a  double  hair  tuft,  sometimes 
inconspicuous  but  sometimes  large  and  erectile,  on  the  sternum  of  the  eighth  segment;  it  is  not 
known  whether  it  plays  any  part  in  courtship.  The  tergum  of  the  eighth  segment  is  unusually  long 
to  accommodate  the  peculiar  genitalia,  and  bears  a  long  apophysis  at  its  proximo-ventral  edge. 
The  wing  venation  shows  a  high  degree  of  individual  variation,  and  does  not  often  provide  good 
characters  for  separating  genera  and  species.  The  forewing  always  has  11  veins  (vein  R4 
missing),  and  veins  Sc  and  RI  are  separate  throughout  their  lengths.  In  the  males  of  most  species 
the  basal  portion  of  vein  M3  is  slightly  swollen  and  devoid  of  normal  cover  scales,  but  bears  small 
to  very  small  specialised  (?scent)  scales.  The  hindwing  has  a  well-developed  humeral  vein  in 
Megalopalpus  and  Lontalius,  but  in  the  other  genera  it  is  only  weakly  developed  or  absent.  The 
male  genitalia  are  highly  characteristic.  The  uncus  and  tegumen  are  in  the  form  of  enormous 
paired  plates,  which  are  attached  to  the  vinculum  only  narrowly  in  the  dorsal  region,  so  that  they 
are  capable  of  considerable  freedom  of  movement .  Articulating  brachia  are  always  present .  The 
vinculum  bears  on  either  side  two  more  or  less  triangular  processes,  one  directed  proximad  and 
the  other  distad.  The  latter  process  is  comparatively  weakly  sclerotised  and  overlies  the  sides  of 
the  tegumen.  The  valvae  are  small,  with  the  outer  dorsal  portion  (sometimes  referred  to  as  the 
ampulla)  bearing  a  dense  hair  fringe.  The  juxta  is  present  as  a  furca  whose  arms  are  united  by  a 
band  above  the  phallus  just  distad  of  the  ductus.  Eliot  (1973:  386)  incorrectly  called  this  band  a 
form  of  transtilla.  The  female  genitalia  have  not  been  investigated. 

The  early  stages  are  very  imperfectly  known.  According  to  Doherty  (1889)  the  eggs  of  Miletus 
and  Allotinus  are  much  flattened  and  disc-like,  but  in  Logania  are  stouter  and  scarcely  more  than 
twice  as  wide  as  high.  They  bear  between  two  and  five  lateral  carinae  which  are  either  simple  or 
broken  into  short  teeth  placed  one  above  the  other,  giving  the  appearance  of  a  cogged  wheel. 
The  larvae,  so  far  as  known,  are  wholly  aphytophagous,  feeding  on  Homoptera.  They  are  more 
or  less  cylindrical  and  have  a  particularly  thick  cuticle,  and  apparently  lack  the  'honey  gland'  on 
the  seventh  and  paired  eversible  tubercles  on  the  eighth  abdominal  segments  which  are  present 
in  the  majority  of  Lycaenidae.  Their  relationship  with  the  ants  attending  the  Homoptera 
appears  to  be  one  of  neutrality,  and  from  this  it  appears  at  least  possible  that  the  larvae  may  be 
furnished  with  small  glands  on  a  number  of  segments,  as  in  many  other  Lycaenidae  (Cottrell, 
1984),  and  that  these  secrete  some  substance  which  inhibits  ant  aggression.  In  one  species, 
Miletus  boisduvali,  the  larva  pupates  inside  ants'  nests  and  the  pupa  has  attractant  glands  and  is 
attended  by  ants;  the  emerging  adult  is  clothed  with  fugitive  scales,  as  in  Liphyra  brassolis, 
which  confuse  attacking  ants  (Roepke,  1918).  However,  in  M .  chinensis,  the  larva  pupates  in  the 
open  and  is  attached  by  the  cremaster  with  or  without  a  weak  girdle  (Kershaw,  1905) ,  and  in  this 
species,  as  well  as  in  Allotinus  subviolaceus,  there  is  no  evidence  to  suggest  that  fugitive  scales 
are  present  in  the  adult  (Piepers  &  Snellen,  1918).  The  adults  feed  on  the  excretions  of 
Homoptera  and  do  not  visit  flowers. 

The  tribe  comprises  four  Oriental  genera  and  one  African  genus  (the  latter  erroneously 
described  from  Borneo).  The  Oriental  genera,  totalling  69  mainly  Sundanian  species,  have  been 


THE  MILETINI  7 

subjected  to  a  full  taxonomic  revision.  No  comparable  attempt  has  been  made  to  revise  the  four 
species  of  Megalopalpus ,  but  a  review  of  the  genus,  based  on  a  draft  kindly  prepared  by  Mr  R.  I. 
Vane-Wright  and  Mrs  S.  M.  North,  is  included. 

In  the  keys  and  descriptions  which  follow,  the  system  of  veins  and  spaces  is  as  in  Fig.  49  (p.  74). 

Key  to  the  genera  of  Miletini 

1  Tarsi  with  first  segment  more  or  less  cylindrical 2 

-  Tarsi  with  first  segment  flattened  and  blade-like 4 

2  Tibiae  swollen  or  incrassate 3 

-  Tibiae  not  swollen;  legs  long,  thin,  cylindrical ALLOTINUS  (p.    7) 

3  Hindwing  without  a  humeral  vein.  Legs  comparatively  short LOGANIA   (p.  57) 

-  Hindwing  with  a  humeral  vein .  Legs  long  and  thin LONTALIUS  (p.  74) 

4  Hindwing  without  a  humeral  vein.  Oriental MILETUS  (p.  75) 

Hindwing  with  a  humeral  vein.  African MEGALOPALPUS  (p.  84) 

Genus  ALLOTINUS  C.  &  R.  Felder 

Allotinus  C.  &  R.  Felder,  [1865]:  285.  Type-species:  Allotinus fallax  C.  &  R.  Felder,  [1865]:  285,  pi.  35,  fig. 
24,  by  designation  of  Scudder,  1875:  107.  Gender  masculine. 

Legs  long,  thin,  cylindrical.  Labial  palpus  with  third  segment  longer  than  half  second  segment.  The  wing 
venation  shows  a  high  degree  of  infra-specific  individual  variation.  Hindwing  cilia  elongated  into  short  tufts 
at  vein  endings  in  the  male,  the  margin  more  or  less  crenulate  and  tufted  in  the  female.  Underside 
characteristic,  white  to  pale  buff,  densely  striated  with  small  striae  or  spots,  with  heavier  spots  arranged  in 
the  usual  lycaenid  pattern.  Forewing  usually  with  very  small  white  costal  flecks  at  ends  of  veins  Sc,  R\,  R2, 
/?3  and,  in  one  species-group,  R5  also. 

The  genus  ranges  from  north  India  to  Sundaland,  the  Philippines  and  Sulawesi. 

Fruhstorfer  (1913-15;  1916)  divided  the  genus  into  two  subgenera:  Allotinus  for  species  allegedly 
without  a  sex  stripe  in  the  male  and  Paragerydus  for  species  with  a  sex  stripe.  I  maintain  these  subgenera, 
though  not  in  the  arrangement  adopted  by  Fruhstorfer,  and  add  a  third  subgenus. 

Key  to  the  subgenera  of  Allotinus 

1  cf  valva  ending  in  an  apical  point;  abdominal  hair  tufts  poorly  developed,  not  protruding  except 

when  genitalia  are  extruded.  Hindwing  usually  with  a  weak  humeral  vein 2 

-  Cf  valva  with  costa  abruptly  truncate  (except  in  A .  davidis) ,  but  ending  in  a  ventral ,  more  or  less 

pointed  process;  abdominal  hair  tufts  prominent.  Hindwing  without  a  humeral  vein 

PARAGERYDUS  (p.  30) 

2  Antenna  barely  longer  than  half  the  forewing  costa,  with  less  than  50  segments,  cf  forewing  with 

vein  A/3  not,  or  only  very  briefly,  swollen ALLOTINUS  (p.    7) 

-  Antenna  nearly  two-thirds  length  of  forewing  costa,  with  60  or  more  segments,  cf  forewing  vein 

M3  prominently  swollen,  clothed  with  specialised  scales  for  one-third  of  its  length  or  longer 

FABITARAS  (p.  20) 

Subgenus  ALLOTINUS  C.  &  R.  Felder 

The  principal  characters  of  the  subgenus  are  as  given  in  the  key. 

Nineteenth  century  authors  separated  Allotinus  from  Paragerydus  by  the  former's  possession  of  a  short 
upper  discocellular  vein,  but  this  character  does  not  occur  in  A.  (A.)  agnolia,  wherein  veins  R5  and  Ml  have 
a  short  common  stalk.  Typical  species,  in  which  the  males  have  vein  M3  unswollen,  have  vein  /?3  long, 
arising  closer  to  the  cell  apex  than  to  the  wing  apex;  but  in  the  two  species  in  which  the  males  have  vein  A/3 
weakly  swollen,  vein  /?3  is  shorter,  arising  close  to  or  opposite  the  end  of  vein  R2. 

On  the  basis  of  the  male  structure  the  subgenus  can  be  divided  into  two  species-groups:  the/fl/fax-group 
in  which  the  abdomen  is  of  normal  length  for  the  tribe  and  the  uncus/tegumen  plates  are  long  and  narrow; 
and  the  major-group  in  which  the  abdomen  is  much  shorter  and  the  uncus/tegumen  plates  are  more 
rounded. 

The  subgenus  has  a  restricted  distribution  in  Sundaland,  the  Philippines  and  Sulawesi,  and  comprises 
seven  species. 


8  J.   N.   ELIOT 

Key  to  the  species  of  subgenus  Allotinus 

1  cf  abdomen  longer  than  hindwing  dorsum.  Underside  of  hindwing  with  postdiscal  spot  in  space 

6  more  or  less  below  the  spot  in  space  7  and  remote  from  the  spot  in  space  5  (fallax- group) 2 

Cf  abdomen  same  length  as  hindwing  dorsum.  Underside  of  hindwing  with  postdiscal  spot  in 
space  6  more  or  less  equidistant  from  the  spots  in  spaces  7  and  5  (major-group) 6 

2  Upperside  with  white  or  grey-blue  areas 3 

Upperside  plain  brown nicholsi  (p.  16) 

3  Upperside  with  white  areas 4 

Upperside  with  greyish-blue  areas subviolaceus  (p.  14) 

4  cf  upperside  of  hindwing  brown  or  diffusely  sullied  with  white  scales 5 

Cf  upperside  of  hindwing  with  a  clearly  defined  white  band albifasciatus  (p.  13) 

5  Forewing  with  veins  R5  and  MI  just  separate  at  their  origins,  cf  forewing  vein  M3  not  swollen  nor 

clothed  with  specialised  scales.  Underside  with  postdiscal  series  of  spots  clearly  marked 

fallax  (p.    8) 

-  Forewing  with  veins  RI  and  M\  stalked,  cf  forewing  vein  M3  briefly  swollen  and  clothed  with 

specialised  scales.  Underside  with  postdiscal  series  not  apparent  on  forewing  and  barely 
discernible  on  hindwing agnolia  (p.  15) 

6  Smaller,  forewing  13-5-19-0  mm.  cf  upperside  of  forewing  all  brown  or  with  a  small  white  or 

whitish  spot  comprising  at  most  a  small  sullied  area  at  base  of  space  2  and  a  larger  white  area 
up  to  3-0  mm  wide  in  space  Ib.  $  with  a  larger  white  area  filling  basal  third  of  space  2  and  about 
half  of  space  Ib,  but  not  reaching  vein  Cu\  nor  entering  the  cell;  white  patch  may  be  reduced, 
sullied  or  absent major  (p.  17) 

-  Larger,  forewing  20-0-21-0  mm.  cf  upperside  of  forewing  with  a  white  patch  filling  basal  third  of 

space  2  and  about  half  of  space  Ib  which  is  basally  grey.  $  with  white  patch  larger,  extending 
above  vein  Cu\  and  into  lower  edge  of  cell maximus  (p.  19) 

Allotinus  (Allotinus)  fallax  C.  &  R.  Felder 

(Figs  1-3,  cf  genitalia) 
Allotinus  fallax  C.  &  R.  Felder,  [1865]:  285,  partim. 

The  Felders  confused  two  distinct  species  under  this  name.  In  pi.  35,  fig.  24  they  figured  as  a  male  what  is 
currently  treated  as  the  female  of  A.  fallax,  and  at  figs  25,  26  as  the  female  a  different  species  which  I 
describe  later  as  the  female  of  A.  (Paragerydus)  albatus  mendax.  Semper  (1889: 163)  correctly  pointed  out 
that  the  Felders'  'male'  was  in  fact  a  female.  However,  he  incorrectly  identified  their  female  as  a  male 


Fig.  1    Allotinus  (Allotinus)  fallax  fallax  C.  &  R.  Felder;  Luzon.  Male  genitalia.  Above,  lateral  view  of 
right  half  of  armature ;  below,  dorsal  view  of  phallus. 


THE  MILETINI  9 

variety  of  fallax,  having  a  white  area  on  the  upperside  of  the  hindwing  such  as  he  had  seen  only  in  a  few 
examples  from  Luzon;  these  may  have  been  either  males  or  females  ofmendax.  Apart  from  these  supposed 
variants  from  Luzon  he  correctly  stated  that  the  male  of  fallax  has  a  plain  brown  hindwing  in  the 
Philippines. 

The  species  is  distinguished  by  lacking  a  swelling  of  vein  A/3  in  the  male  and  by  the  presence  of  a  white 
patch  on  the  upperside  of  the  dark  brown  forewing.  The  hindwing  of  the  male  is  brown,  except  in  Borneo 
where  it  is  usually  dusted  with  white  scales.  Except  in  Sumatra  and  the  Malay  Peninsula,  where  the 
hindwing  is  brown,  the  female  has  a  white  discal  area  (rarely  vestigial  or  absent)  which  shows  a  high  degree 
of  individual  variation,  so  that  it  is  an  unreliable  character  by  which  to  distinguish  subspecies. 

On  the  basis  of  the  male  genitalia  A.  fallax  can  be  divided  into  three  subspecies-groups. 

Type  (a)  (Fig.  1):  valva  with  short  apical  hook  and  ventro-distal  edge  not  strongly  exarcate;  phallus 
comparatively  short,  broad  and  abruptly  tapered  at  distal  end;  Philippine. 

Type  (b)  (Fig.  2):  valva  with  short  apical  hook  and  ventro-distal  edge  strongly  exarcate;  phallus 
comparatively  long  and  thin  and  gradually  tapered  at  distal  end;  Philippine. 


Fig.  2    Allotinus  (Allotinus)  fallax  aphacus  Fruhstorfer ;  Mindanao.  Male  genitalia. 

Type  (c)  (Fig.  3):  valva  with  long  apical  hook;  phallus  like  type  (c),  but  still  slimmer;  Sundanian. 

So  far  as  my  experience  goes  type  (a)  occurs  exclusively  in  Luzon,  Masbate,  Bohol,  Samar  and  Leyte, 
and  type  (b)  exclusively  in  Mindanao  and  Bazilan.  Both  occur  in  Mindoro,  each  type  having  a  distinctive 
phenotype.  I  have  not  been  able  to  dissect  males  from  Cebu,  Panaon,  Camiguin  de  Mindanao  and  the  Sulu 
Is. ,  whence  Fruhstorfer  named  subspecies,  but  I  think  it  certain  that  type  (a)  will  be  found  in  Cebu  and  type 
(b)  in  Camiguin  de  Mindanao  and  Sulu  Is. ,  whilst  in  Panaon  either  might  occur.  The  occurrence  of  type  (a) 
in  Bohol,  Samar  and  Leyte  is  rather  unexpected;  usually  butterflies  from  these  islands  are  associated  with 
Mindanao  rather  than  with  Luzon.  The  dichotomy  in  Mindoro  can  be  paralleled  in  the  lycaenopsid 
Acytolepis puspa  which  occurs  in  Mindoro  in  subsp.  cagaya  and  the  very  different-looking  subsp.  bazilana 
in  equality  and  without  evidence  of  intergradation  (Eliot  &  Kawazoe,  1983:  183).  Conceivably  types  (a) 
and  (b)  may  represent  distinct  species  which  evolved  in  the  northern  and  southern  groups  of  islands 
respectively  and  which  may  now  have  a  wider  overlap  in  distribution  than  is  at  present  known;  otherwise  it 
is  difficult  to  see  how  type  (b)  could  have  reached  Mindoro  without  also  becoming  established  in  the 
intervening  islands. 

A.  fallax  flies  at  low  to  moderate  elevations,  and  is  common  and  widespread  in  the  Philippines,  but  rare 
in  Sundaland.  It  has  not  been  found  in  Palawan,  but  is  likely  to  occur  there. 


Key  to  the  subspecies  of  A.  (A.)  fallax 
1     cf  genitalia  of  type  (c) .  Sundanian 


-    cf  genitalia  of  types  (a)  or  (b).  Philippine 

2     cf  genitalia  of  type  (b).  $  upperside  of  hindwing  with  white  area  normally  confined  to  basal 
two-thirds  of  spaces  4  and  5 ,  but  may  be  vestigial  or  obsolete 


10  J.   N.  ELIOT 

Cf  genitalia  of  type  (a).  $  upperside  of  hindwing  with  white  area  normally  extending  below  vein 
A/3,  often  as  far  as  vein  A! fallax  fallax  (p.  10) 

3  cf  upperside  of  forewing  with  white  patch  reaching  vein  Cu\ 5 

Cf  upperside  of  forewing  with  white  patch  usually  not  above  mid-space  2;  if  reaching  vein  Cui, 

upper  part  narrow  and  sullied 4 

4  cf  white  patch  comparatively  large,  reaching  into  space  2.  Underside  of  forewing  with  central 

area  above  dorsum  sparsely  striated fallax  aphacus  (p.  11) 

Cf  sullied  white  patch  smaller,  usually  confined  to  space  Ib.  Underside  of  forewing  with  central 
area  above  dorsum  as  densely  striated  as  rest  of  wing fallax  eryximachus  (p.  11) 

5  Underside  ground  colour  pale  buff.  $  upperside  of  hindwing  with  a  small  white  patch 

fallax  dotion  (p.  12) 
Underside  ground  colour  whitish  buff,  with  pale  markings.  $  hindwing  white  patch  obsolete 

fallax  tymphrestus  (p.  12) 

6  Upperside  of  hindwing  with  a  dusting  of  white  scales  usually  present  in  cf  and  always  present  in 

$ fallaxaudax  (p.  12) 

Upperside  of  hindwing  plain  brown fallax  apus  (p.  12) 

Allotinus  (Allotinus)  fallax  fallax  C.  &  R.  Felder 
(Fig.  1,  cf  genitalia) 

Allotinus  fallax  C.  &  R.  Felder,  [1865]:  285,  partim  'cf ',  recte  $  [nee  $ ,  pi.  35,  fig.  24  'cf'];  Semper,  1889: 
163,  partim,  pi.  31,  figs  23cf,  26$.  LECTOTYPE  $,  PHILIPPINES:  Luzon  (BMNH),  here  designated 
[examined]. 

Allotinus  fallax  fallax  Felder;  Fruhstorfer,  1913:  343;  1916:  809. 

Allotinus  fallax  sabazus  Fruhstorfer,  1913:  343;  1916:  809.  LECTOTYPE  cf,  PHILIPPINES:  Bohol  [not 
located] .  Syn.  n. 

Allotinus  fallax  zaradrus  Fruhstorfer,  1916:  809.  Syntypes,  PHILIPPINES:  Cebu  [not  located]. 

In  the  male  the  white  patch  on  the  forewing  almost  always  extends  across  vein  Cu\  into  space  3.  In  the 
female  the  white  patch  on  the  hindwing  is  variable  in  extent,  but  almost  always  extends  dorsad  into  space  3 
and  often  as  far  as  space  Ib.  The  ground  colour  of  the  underside  is  pale  buff,  richer  in  the  male  than  in  the 
female. 

The  Felders  did  not  specify  a  type.  Their  'male',  as  pointed  out  by  Semper  (1889: 163),  is  a  female,  while 
their  female  belongs  to  another  species  described  later  as  A.  (Paragerydus)  albatus  mendax.  In  BMNH 
there  is  a  male  ex  Felder  coll.  bearing  a  BMNH  red  type  label,  but  as  the  original  figure  and  description 
does  not  apply  to  this  sex  it  seems  better  to  reject  this  unpublished  type  selection  and  to  designate  as 
lectotype  a  female  in  BMNH  labelled  /79/Luzon  Lorquin  [round  blue]/Felder  Coll. /Rothschild  Bequest 
1939-17. 

When  naming  sabazus,  Fruhstorfer,  evidently  referring  to  the  male,  stated  that  the  white  patch  on  the 
forewing  was  reduced  compared  with  subsp.  fallax  and  that  the  underside  was  darker  and  more  closely 
striated.  The  male  from  Bohol  figured  by  Semper  has  the  white  patch  entirely  below  vein  C«i,  as  in 
occasional  Luzon  examples,  but  in  a  series  of  both  sexes  from  Bohol  in  coll.  Treadaway  the  males  have  the 
white  patch  extending  above  vein  Cu\  and  in  neither  sex  can  I  detect  any  consistent  differences  from  Luzon 
fallax.  I  therefore  consider  that  sabazus  cannot  be  maintained  as  a  valid  subspecies.  There  is  in  SM  a  female 
labelled  /Bohol/Coll.  C.  Semper/2 11/Typus  [red]/,  but  in  my  view  it  cannot  be  accepted  as  the  type  of 
sabazus  since  Fruhstorfer's  description  did  not  apply  to  this  sex.  I  therefore  designate  as  lectotype  the  male 
figured  by  Semper  (pi.  31,  fig.  23),  which  unfortunately  has  not  been  recognised  in  SM. 

Semper  recorded  fallax  from  Cebu,  but  did  not  figure  it.  It  is  not  clear  whether  Fruhstorfer  (1916)  was 
merely  attaching  a  name  to  Semper's  record  or  actually  saw  specimens  from  Cebu.  No  examples  from  Cebu 
can  now  be  found  in  SM,  but  according  to  Treadaway  (pers.  comm.)  it  is  thought  that  some  may  possibly 
exist  but  be  temporarily  mislaid.  The  only  specimen  that  I  have  seen  from  Cebu  is  a  female  in  coll. 
Treadaway.  This  has  the  underside  ground  colour  somewhat  whiter  than  usual,  but  the  difference  from 
Luzon  females  seems  too  slight  to  justify  retaining  zaradrus  as  a  valid  subspecies,  and  I  therefore 
provisionally  synonymise  it  with /a/tax. 

Examples  from  Mindoro  and  Sibuyan  Is.  do  not  differ  from  normal  fallax,  but  two  males  from  Masbate 
in  coll.  Treadaway  have  a  more  whitish  underside,  and  a  female  from  Panay  is  almost  pure  white  and 
relatively  weakly  striated  below  and  has  an  exceptionally  large  white  patch  on  the  upperside  of  the 
hindwing.  I  provisionally  place  these  under  fallax . 

Males  from  Samar  and  Leyte  seem  never  to  have  the  forewing  white  patch  extending  above  vein  Cui  and 


THE  MILETINI  11 

in  both  sexes  the  underside  ground  colour  is  darker  buff  than  in  examples  from  Luzon  and  Bohol.  These 
possibly  constitute  a  distinct  subspecies  to  which  Fruhstorfer's  name  artinus,  given  by  him  to  the  population 
in  Panaon,  may  apply.  Unfortunately  I  have  seen  no  males  from  Panaon  and  Semper's  figure  of  a  Panaon 
male  (pi.  31,  fig.  24)  shows  only  the  underside,  from  which  it  is  not  possible  to  establish  whether  or  not  it 
resembles  Samar  and  Leyte  examples.  The  single  female  from  Panaon  which  I  have  seen  does  not  differ 
from  Mindanao  females,  and  I  therefore  provisionally  synonymise  artinus  with  aphacus. 

DISTRIBUTION.  Luzon,  Mindoro,  Sibuyan,  Cebu,  Bohol.  In  slightly  different  forms  also  in  Masbate,  Panay, 
Samar  and  Leyte.  I  have  seen  no  examples  from  Negros,  but  it  must  occur  there  as  well  as  in  other  islands. 

Allotinus  (Allotinus)  fallax  aphacus  Fruhstorfer 
(Fig.  2,  cf  genitalia) 

Allotinus  fallax  C.  &  R.  Felder;  Semper,  1889: 163,  partim,  pi.  31,  figs  24  cf ,  25  $ . 

Allotinus  fallax  aphacus  Fruhstorfer,  1913:  343;  1916:  809.  Holotype  $,  PHILIPPINES:  Camiguin  de 

Mindanao  [not  located]. 
Allotinus  fallax  ancius  Fruhstorfer,  1913:  343;  1916:  809.  LECTOTYPE  cf ,  PHILIPPINES:  Mindanao 

(BMNH),  here  designated  [examined].  Syn.  n. 
Allotinus  fallax  artinus  Fruhstorfer,  1916:  809.  LECTOTYPE  $,  PHILIPPINES:  Panaon  (SM),  here 

designated  [examined]. 

On  the  upperside  the  forewing  white  patch  of  the  male  is  barely  half  as  large  as  in  subsp.  fallax,  seldom  fully 
crossing  space  2,  and  when  it  does  so  the  upper  part  is  sullied  with  brown  scales.  The  female  has  the  white 
patch  on  the  hindwing  almost  always  restricted  to  the  basal  two-thirds  of  spaces  4  and  5,  and  often  it  is 
almost  obsolete.  On  the  underside  of  the  forewing  there  is  an  almost  unstriated  central  area  above  the 
dorsum.  The  male  genitalia  are  type  (b). 

Fruhstorfer  named  aphacus  from  Semper's  record  and  figure  25  of  a  female  from  Camiguin  de 
Mindanao.  The  original  of  this  figure,  which  I  have  not  been  able  to  locate,  is  therefore  automatically  the 
holotype.  I  have  seen  a  single  male  labelled  /Cam.  de  Mind./211/Coll.  C.  Semper/  which  does  not  differ 
from  Mindanao  males. 

I  designate  as  lectotype  of  ancius  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Mindanao/Mindanao  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/Allotinus  fallax  ancius  [in  Corbet's 
hand]/.  A  female  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/Mindanao/Mindanao  Fruhstorfer/Fruh- 
storfer Coll.  B.M.  1933-131/fallax  ancius  Fruhst.  [in  Fruhstorfer's  hand]/  becomes  a  paralectotype. 

I  designate  as  lectotype  of  artinus  a  female  in  SM  labelled  /Coll.  C.  Semper/Panaon/211/Typus  [red]/, 
which  does  not  differ  from  Mindanao  examples. 

DISTRIBUTION.  Mindanao,  Camiguin  de  Mindanao,  Panaon.  I  also  provisionally  place  under  this  subspecies 
a  female  in  BMNH  from  Talaud  I. 

Allotinus  (Allotinus)  fallax  eryximachus  Fruhstorfer 

Allotinus  fallax  eryximachus  Fruhstorfer,  1913:  343;  1916:  809.  LECTOTYPE  cf ,  PHILIPPINES:  Mindoro 
(BMNH),  here  designated  [examined]. 

This  is  the  darkest  subspecies.  As  pointed  out  by  Fruhstorfer  in  his  original  description,  the  forewing  white 
patch  in  the  male  is  even  more  insignificant  (unbedeutenderen)  than  in  subsp.  aphacus,  being  always  more 
or  less  sullied  with  brown  scales  and  sometimes  restricted  to  a  mere  streak  below  vein  Cu2.  On  the 
underside  the  central  area  above  the  forewing  dorsum,  which  is  not,  or  is  comparatively  weakly,  striated  in 
the  other  subspecies,  is  striated  all  over.  The  male  genitalia  are  type  (b). 

Fruhstorfer  stated  that  the  type  was  in  BMNH.  However,  there  are  no  males  agreeing  with  his 
description  labelled  as  types,  but  a  pair  of  subsp.  fallax  have  been  placed  in  the  type  drawer  above  the  name 
eryximachus.  The  male,  with  a  large  white  forewing  patch,  is  labelled  /Mindoro  Philippine  Is.  Dr. 
Platen/Godman-Salvin  Coll.  1908-168/A.  fallax  107  Var.  ?/,  and  the  female  is  labelled  like  the  male  but 
with  an  additional  label  /Allotinus  'cf '  fallax  Felder/.  Agrees  with  figure  of  type  F.  A.  H.  8.  x.  09/.  Neither 
specimen  bears  a  BMNH  red  type  label,  presumably  because  they  cannot  be  reconciled  with  Fruhstorfer's 
description  of  eryximachus.  Although  Fruhstorfer  may  have  seen  this  pair  in  BMNH  they  are  not  to  be 
considered  as  types,  and  I  designate  as  lectotype  a  male  from  a  series  which  Fruhstorfer  may  have  seen  at 
Tring  and  which  he  may  have  confused  with  what  he  saw  in  BMNH;  it  is  labelled  /Mindoro  Platen/ 
Rothschild  Bequest  1939-1/.  It  is  interesting  that  Platen  should  have  caught  both  subspecies  in  Mindoro. 

DISTRIBUTION.  Mindoro,  where  it  appears  to  outnumber  sympatric  subsp.  fallax. 


12  J.   N.  ELIOT 

Allotinus  (Allotinus)  fallax  dotion  Fruhstorfer 

Allotinus  fallox  dotion  Fruhstorfer,  1913:  343;  1916:  809,  pi.  141h  cf  $  as  'dolion\  LECTOTYPE  cf, 
PHILIPPINES:  Bazilan  (BMNH),  here  designated  [examined]. 

On  the  upperside  both  sexes  resemble  subsp.  fallax,  with  the  white  patch  on  the  forewing  of  the  male 
unsullied  and  reaching  vein  €1*2  broadly.  On  the  underside  the  ground  colour  is  a  paler,  more  whitish  buff 
than  in  subspp.  sabazus  and  aphacus.  The  male  genitalia  are  type  b. 

I  designate  as  lectotype  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/Philippinen 
Bazilan  II-III.  98  Doherty  ex  coll.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/fallax  dotion  Fruhst.  [in 
Fruhstorfer's  hand]/.  A  female  labelled  similarly,  except  that  the  final  label  reads  /dotion  [in  Fruhstorfer's 
hand]/,  is  labelled  paralectotype. 

DISTRIBUTION.  Bazilan. 

Allotinus  (Allotinus)  fallax  tymphrestus  Fruhstorfer 

Allotinus  fallax  tymphrestus  Fruhstorfer,  1916:  809.  LECTOTYPE  cf,  PHILIPPINES:  Sulu  Is.  (SM),  here 
designated  [examined] . 

I  have  seen  only  the  original  pair  from  which  Fruhstorfer  described  the  subspecies.  The  male  exactly 
resembles  subsp.  dotion  on  the  upperside,  but  on  the  underside  the  ground  colour  is  slightly  more  whitish 
and  the  markings  less  well-defined.  The  female  lacks  the  usual  white  patch  on  the  upperside  of  the 
hindwing,  but  it  may  be  that  a  white  patch  is  sometimes  present.  The  subspecies  is  doubtfully  separate  from 
subsp.  dotion. 

I  designate  as  lectotype  the  male  in  SM  labelled  /Coll.  C.  Semper/Sulu  Inseln  Meyerink/211/Typus 
[red]/.  The  female  is  labelled  paralectotype. 

DISTRIBUTION.  Sulu  Is. 

Allotinus  (Allotinus)  fallax audaxH.  H.  Druce 

Allotinus  audox  H.  H.  Druce,  1895:  564,  pi.  31,  figs  11  cf ,  12  $>.  LECTOTYPE  cf ,  BORNEO:  Mt  Kina  Balu 

(BMNH),  here  designated  [examined]. 
Allotinus  fallax  audax  Druce;  Fruhstorfer,  1913:  343;  1916:  809;  Corbet,  1939ft:  66. 

In  both  sexes  the  forewing  white  patch  is  more  extensive  than  in  the  Philippine  subspecies,  almost  reaching 
the  wing  base  in  space  Ib  where  it  is  dusted  with  grey  scales.  The  hindwing  is  more  or  less  lightly  dusted  with 
white  scales  in  the  male,  but  heavily  dusted  over  most  of  the  wing  in  females. 

The  subspecies  was  described  from  an  unspecified  number  of  specimens  in  coll.  H.  H.  Druce  and 
Staudinger.  I  designate  as  lectotype  a  male  in  BMNH  labelled  /Kina  Balu  Waterstr.  A.  audax  co-type 
H.  H.  Druce/ex  coll.  Hamilton  Druce  1919/Type  PT  [green]/Joicey  Bequest  Brit.  Mus.  1934-120/.  A 
female  labelled  $  co-type,  but  otherwise  as  the  male,  is  a  paralectotype. 

DISTRIBUTION.  Borneo;  apparently  only  known  from  Mt  Kina  Balu. 

Allotinus  (Allotinus)  fallax  apus  de  Niceville 
(Fig.  3,  cf  genitalia) 

Allotinus  apus  de  Niceville,  1895:  27,  pi.  S,  fig.  17  $;  Fruhstorfer;  1913:  343;  1916:  809.  Syntypes  $, 

SUMATRA  (?  ZSI). 
Allotinus  fallax  apus  de  Niceville;  Corbet,  1939ft:  66;  Eliot,  1962:  218;  1978:  240;  1980: 145,  figs  7  cf ,  14  cf 

genitalia;  Fleming,  1975;  22,  pi.  50,  fig.  L50  cf . 
Allotinus  fallax  michaelis  Eliot,  1959:  376,  partim  cf  nee  $ ,  pi.  10,  fig.  8  cf .  Holotype  cf ,  WEST  MALAYSIA 

(BMNH)  [examined].  [Synonymised  by  Eliot,  1962:  218.] 

Differs  from  subsp.  audax  in  the  plain  brown  hindwing  in  both  sexes. 

A.  apus  was  described  from  two  females  from  north-east  Sumatra,  which  should  be  in  ZSI.  There  is  in 
BMNH  a  female  labelled  /Type  [red]/Battak  Mts.  N.E.  Sum.  11.  94  Dr  Martin/ Allotinus  apus  de  Niceville 
and,  [on  reverse] ,  Collect  Dr.  Martin/Rothschild  Bequest  B.M.  1939-1/.  As  there  is  no  evidence  to  suggest 
that  this  female  was  one  of  de  Niceville's  original  two  I  reject  it  as  a  type. 

DISTRIBUTION.  Sumatra  and  West  Malaysia,  at  elevations  of  about  500  m  upwards. 


THE  MILETINI 


13 


Fig.  3    Allotinus  (Allotinus)  fallax  apus  de  Niceville;  Malay  Peninsula.  Male  genitalia. 


Allotinus  (Allotinus)  albifasciatus  Eliot 

(Fig.  4,  cf  genitalia) 

[Allotinus fallax  michaelis  Eliot,  1959:  376  (partim,  $  nee  cf),  pi.  10,  fig.  9  <j>.  Misidentification.] 
[Allotinus  fallax  audax  H.  H.  Druce  sensu  Eliot,  1978:  241.  Misidentification.] 

Allotinus  albifasciatus  Eliot,  1980:  143,  figs  5  d",  6  <j>,  14  cf  genitalia.  Holotype  cf,  SUMATRA  (BMNH) 
[examined]. 

The  species  is  distinctive  in  possessing  a  clear  white  band  crossing  the  hindwing  in  both  sexes.  On  the 
underside  the  ground  colour  is  off-white,  with  the  postdiscal  markings  larger  and  more  blotchy  than  in  A. 
fallax. 

DISTRIBUTION.  The  species  is  extremely  rare,  and  has  so  far  only  been  found  in  the  Malay  Peninsula  and 
Sumatra  at  elevations  of  about  1000  m. 


Fig.  4    Allotinus  (Allotinus)  albifasciatus  Eliot;  Sumatra.  Male  genitalia. 


14  J.  N.  ELIOT 

Allotinus  (Allotinus)subvioIaceusC.  &  R.  Felder 

(Fig.  5,  cf  genitalia) 
Allotinus subviolaceus  C.  &  R.  Felder,  [1865]:  286. 

The  greyish  blue  areas  on  the  upperside  render  this  species  unmistakable.  As  in  the  white  areas  of  A.  fallax 
these  blue  areas  are  individually  variable  in  extent,  especially  on  the  hindwing  where  they  may  be  absent  or 
cover  almost  the  whole  wing.  On  average  they  are  more  extensive  on  both  wings  in  the  female. 

The  species  has  the  widest  distribution  in  the  subgenus,  occurring  from  Assam  to  Java  and  the 
Philippines. 

Key  to  the  subspecies  of  A.  subviolaceus 

1     Upperside  greyish  blue.  Upperside  of  forewing  in  cf  with  space  3  all  or  almost  all  black 

subviolaceus  subviolaceus  (p.  14) 

Upperside  pale  grey,  with  only  a  slight  blue  tint ,  becoming  whitish  in  lower  half  of  forewing  disc. 
Upperside  of  forewing  in  cf  with  basal  half  of  space  3  and  base  of  space  4  bluish  grey 

subviolaceus  mirus  (p.  15) 


Allotinus  (Allotinus)  subviolaceus  subviolaceus  C.  &  R.  Felder 

(Fig.  5  cf  genitalia) 

Allotinus  subviolaceus  C.  &  R.  Felder,  [1865]:  286,  pi.  35,  figs  27, 28  cf ;  Piepers  &  Snellen,  1918: 17,  pi.  20, 
figs  20a  cf ,  20b  $ ,  20c  larva.  LECTOTYPE  cf ,  JAVA  (BMNH),  here  designated  [examined]. 

Allotinus  alkamah  Distant,  18860:  452,  pi.  44,  'cf '  recte  <j>.  Holotype  <j>,  WEST  MALAYSIA  (not  located). 
Syn.  n. 

Allotinus  subviolescens  [sic]  Felder;  Swinhoe,  1910:  196,  pi.  616,  figs  1,  la  $,  Ib  cf. 

Allotinus  subviolaceus  alkamah  Distant;  Fruhstorfer,  1913:  342;  1916:  808;  Corbet,  1939a:  66;  Eliot,  1978: 
240;  Fleming,  1975:  pi.  57,  fig.  L37  $. 

Allotinus  subviolaceus  subviolaceus  C.  &  R.  Felder;  Fruhstorfer,  1913:  342;  1916:  808,  pi.  141g  £. 

Allotinus  subviolaceus  manychus  Fruhstorfer,  1913:  342;  1916:  808;  Evans,  1932:  212;  Cantlie,  1967:  28. 
LECTOTYPE  <j>,  BURMA  (BMNH),  here  designated  [examined].  Syn.  n. 

Allotinus  subviolaceus  kallikrates  Fruhstorfer,  1913:  342;  1916:  808.  LECTOTYPE  cf,  PHILIPPINES: 
Mindanao  (BMNH),  here  designated  [examined].  Syn.  n. 

Allotinus  subviolaceus  silarus  Fruhstorfer,  1916:  808.  LECTOTYPE  cf ,  BORNEO  (BMNH),  here  desig- 
nated [examined].  Syn.  n. 

Males  have  a  broad  black  border  on  the  forewing  which  often  fills  the  whole  of  space  3  and  the  cell,  but  in 
some  examples  the  blue  may  enter  the  lower  half  of  space  3  and  fill  the  lower  two-thirds  of  the  cell.  Usually 
the  hindwing  has  only  a  few  blue  scales. 

In  the  female  the  forewing  border  is  much  narrower,  and  the  hindwing  varies  individually  from  wholly 
fuscous  to  nearly  all  blue. 

In  Palawan  the  greyish  blue  colour  tends  to  be  paler  in  both  sexes  than  in  other  areas,  and  this  does  not 
appear  to  be  due  to  season,  as  in  the  still  paler  examples  flying  in  Burma  and  Thailand  during  the  dry 
season.  In  addition  the  forewing  border  tends  to  be  wider  in  females,  measuring  an  average  of  2-5  mm  at 
vein  Cu2  compared  with  an  average  of  2-0  mm  in  normal  examples.  These  differences  seem  hardly  sufficient 
to  justify  erecting  another  subspecies.  Females  from  Borneo  and  Bangka  also  have,  on  average,  the 
forewing  border  similar  to  Palawan  examples. 

The  Felders  described  subviolaceus  only  from  the  male,  but  did  not  designate  a  type.  I  therefore 
designate  as  lectotype  a  male  in  BMNH  labelled  /Type  [red]/Java  Coll.  .  .  .  [illegible]/[circular  blue]/ 
Allotinus  subviolaceous  Feld. /Felder  COLLN. /subviolaceus  n. /Rothschild  Bequest  B.M.  1939-1/. 

The  holotype  of  alkamah  is  the  'male'  (recte  female)  figured  by  Distant,  which  has  a  fuscous  hindwing.  In 
BMNH  there  is  a  Sumatran  female  labelled  /Type  [red]/alkamah  Dist. /Sumatra  Forbes/Rothschild 
Bequest  B.M.  1939-1/alkamah  Dist.  $  Allotype  [in  script  believed  to  be  by  Corbet]/.  As  the  holotype  is  a 
female  the  allotype  cannot  be  another  female ,  and  I  reject  it  as  such .  Fortunately  Corbet  does  not  appear  to 
have  published  this  type  selection. 

Fruhstorfer  did  not  designate  a  type  of  manychus.  He  had  no  examples  from  Burma  in  his  collection,  but 
said  that  he  had  examined  examples  in  BMNH  from  Pegu  and  Rangoon.  There  is  a  female  in  BMNH 
labelled  /Type  [red]/Moore  Coll.  1908-208  Pegu  Magaree/May  be  taken  as  type  of  Allotinus  subviolaceus 


THE  MILETINI 


15 


Fig.  5    Allotinus  (Allotinus)  subviolaceus  subviolaceus  C.  &  R.  Felder;  Malay  Peninsula.  Male  genitalia. 

manychus  Fruh.  [in  script  believed  to  be  by  Corbet]/.  As  it  is  likely  that  Fruhstorfer  saw  this  specimen 
during  his  visit  to  BMNH I  designate  it  as  lectotype. 

I  designate  as  lectotype  of  kallikrates  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer 
orangej/Mindanao  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/subviolaceus  kallikrates  Fr.  [in  Fruhstor- 
fer's  hand]/.  A  female  labelled  /Type  [red]/Type  [Fruhstorfer  orangej/Mindanao/Mindanao  Fruhstorfer/ 
Fruhstorfer  Coll.  B.M.  1933-131/Allotinus  subviolaceus  kallikrates  Fruh.  [in  Corbet's  hand]/  is  a  paralec- 
totype. 

I  designate  as  lectotype  of  silarus  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Nordborneo  Brunei  Waterstradt  1890/Fruhstorfer  Coll.  B.M.  1933-131/Allotinus  subviolaceus  silarus 
Fruh.  [in  Corbet's  hand]/.  A  female  in  BMNH  labelled  /Type  [red]/W.  Borneo  Sintang  16.  III.  10/ 
Fruhstorfer  Coll.  B.M.  1933-131/subviolaceus  silarus  Fr.  [in  Fruhstorfer's  hand]/  is  labelled  paralectotype. 

DISTRIBUTION.  Manipur;  Burma;  Thailand;  West  Malaysia;  Singapore;  Sumatra;  Bangka  I.;  Java;  Borneo; 
Palawan;  Mindanao. 

Allotinus  (Allotinus)  subviolaceus  mirus  van  Eecke 

Allotinus  subviolaceus  mirus  van  Eecke,  1914: 247;  Fruhstorfer,  1916, 808.  LECTOTYPE  cf ,  SIMEULUE  I. 
(Simalur)  (RNH),  here  designated  [examined]. 

The  subspecies  is  much  paler  than  the  palest  dry  season  examples  from  Burma. 

Above,  both  sexes  are  pale  bluish  grey,  becoming  whitish  on  the  forewing  in  the  middle  of  space  Ib  and 
basal  half  of  space  2.  In  the  male  the  forewing  border  is  inwardly  rather  irregular  and  narrow,  leaving  the 
basal  half  of  space  3 ,  the  base  of  space  4  and  the  lower  three-quarters  of  the  cell  grey ;  it  is  narrowest  at  vein 
Cu2,  where  it  is  just  over  2-0  mm  wide.  The  hindwing  is  mostly  grey,  with  the  discocellular  veins  darkened 
and  with  an  inwardly  diffuse  fuscous  marginal  and  costal  border  2-3  mm  wide.  The  female  differs  as  usual 
in  having  more  extensive  bluish  grey  areas  on  both  wings. 

I  designate  as  lectotype  a  male  in  RNH  labelled  /60.  49/E.  Jacobson  Sinabang.  Sim.  Sum.  7.  1913/ 
Museum  Leiden  Allotinus  subviolaceus  mirus  Det.  R.  v.  Eecke/Type/.  A  further  male  and  a  female 
labelled  /60.  50/  and  /60,  51/  respectively,  but  otherwise  as  the  lectotype,  are  paralectotypes. 

DISTRIBUTION.  Simeulue  I.  (Simalur). 


Allotinus  (Allotinus)  agnolia  sp.  n. 

(Figs  6  cf  genitalia  and  scales;  55  9) 

Cf  forewing  length  12-5  mm.  Forewing  with  apex  rounded,  as  in  females  of  the  genus;  veins  M\  and  R5  with 
a  short  common  stalk  and  vein  R3  shorter  than  in  the  three  preceding  species,  arising  only  a  little  before  end 


16 


J.  N.  ELIOT 


Fig.  6    Allotinus  (Allotinus)  agnolia  sp.  n.:  Sumatra.  Male  genitalia.  Lower  left,  androconial  scale  from 
fore  wing  vein  M3;  below,  normal  cover  scale  from  the  same  area. 


of  vein  R2.  Upperside  dark  brown,  with  basal  quarter  of  forewing  a  paler  grey  brown.  Forewing  with  a 
rhomboidal  white  patch  about  3-5  mm  broad  resting  on  vein  A\  and  reaching  into  lower  angle  of  cell  and 
base  of  space  3.  Basal  quarter  of  vein  A/3  rather  weakly  swollen  and  clothed  with  very  pale  buff  specialised 
scales  about  one-sixth  size  of  white  cover  scales.  Underside  with  a  whitish  area  above  mid-dorsum  of 
forewing ,  but  rest  of  both  wings  pale  buff  very  densely  striated  with  buff-brown  striae ;  usual  postdiscal  and 
submarginal  markings  of  the  genus  small  and  only  made  out  with  difficulty. 

9 .  Apart  from  lacking  the  swelling  of  vein  M3  of  forewing,  similar  in  all  respects  to  male. 

MATERIAL  EXAMINED 

Holotype  cf ,  Sumatra:  'Battak  Mts.,  N.E.  Sum.,  II.  94  (Dr.  Martin)'  (BMNH). 

Paratypes.  Sumatra:  1  $  (allotype),  'VIII.  94',  otherwise  as  holotype  (BMNH);  1  9  'N.O.  Sumatra 
Martin  Coll.  H.  Fruhstorfer',  'XL  94',  'Fruhstorfer  Coll.  B.M.  1933-131'  (BMNH). 

The  specific  name  is  an  anagram  of  Logania,  given  because  the  holotype  was  found  in  BMNH  among  a 
series  of  Logania  distanti  massalia  bearing  a  manuscript  label  (in  an  unknown  hand)  reading  Logania 
massalia  Doherty. 

Allotinus  (Allotinus)  nicholsi  Moulton 

(Figs  7,  cf  genitalia;  56  cf ;  57  $) 
Allotinus  nicholsi  Moulton,  1911:  83. 

The  species  differs  from  all  the  preceding  in  being  reddish  brown  without  white  or  blue  areas.  On  the 
forewing  vein  R3  is  as  short  as  in  Paragerydus,  arising  opposite  the  end  of  vein  R2.  On  the  hindwing  a  weak 
humeral  vein  is  present.  In  the  male,  vein  M3  of  the  forewing  is  weakly  swollen  in  its  basal  quarter  (a  fact 
not  noticed  by  Moulton  and  Corbet) ,  and  the  specialised  scales  are  slightly  smaller  than  those  of  A .  agnolia . 
The  male  genitalia  possess  a  feature  unique  in  the  tribe,  viz.  a  prominent  cornutus  in  the  phallus,  consisting 
of  a  longer  central  and  shorter  outer  bundles  of  minute  spicules. 

The  species  is  known  only  from  Borneo  and  Sumatra,  and  appears  to  be  very  rare.  Provisionally  I 
recognise  two  subspecies,  but  more  material  may  show  that  no  subdivision  is  necessary. 

Key  to  the  subspecies  of  A.  (A.)  nicholsi 

1     Underside  markings  faint  and  comparatively  broad nicholsi  nicholsi  (p.  17) 

-    Underside  markings  narrow,  darker  and  sharply  defined nicholsi  battakanus  (p.  17) 


THE  MILETINI 


17 


Fig.  7    Allotinus  (Allotinus)  nicholsi  battakanus  Fruhstorfer;  Sumatra.  Male  genitalia. 


Allotinus  (Allotinus)  nicholsi  nicholsi  Moulton 

Allotinus  nicholsi  Moulton,  1911:  83;  Fruhstorfer,  1913:  343;  1916:  809;  Corbet,  19396:  66,  fig.  11  cT 

genitalia.  Holotype  cf ,  BORNEO:  Sarawak  (BMNH)  [examined]. 
Allotinus  nicholsi  nicholsi  Moulton;  Eliot,  1967:  71. 

The  subspecies  is  known  only  from  the  male  holotype,  which  is  in  worn  condition.  On  the  forewing  veins 
MI  and  R5  are  just  separate.  On  the  underside  the  markings  are  rather  faint  and  comparatively  wide,  and 
resemble  those  of  A.  subviolaceus,  as  pointed  out  by  Moulton. 

Allotinus  (Allotinus)  nicholsi  battakanus  Fruhstorfer 

(Figs  7  cf  genitalia;  56  cf ;  57  9) 

Allotinus  taros  battakanus  Fruhstorfer,  1913:  370  (partim);  1916:  813,  ?  pi.  141g  $  (partim).  LECTOTYPE 

Cf ,  SUMATRA  (BMNH),  here  designated  [examined]. 
Allotinus  nicholsi  battakanus  Fruhstorfer;  Eliot,  1967:  71. 

The  short  series  of  one  male  and  four  females  in  BMNH  have  veins  M^  and  R5  of  the  forewing  connate ,  and 
on  the  underside  the  markings  are  narrower,  darker  and  more  sharply  defined  than  in  subsp.  nicholsi. 

Although  Fruhstorfer  drew  attention  to  the  absence  of  the  usual  forewing  brand  of  the  male,  part  of  his 
type-series,  including  two  female  syntypes  in  BMNH,  comprise  A.  (Fabitaras)  sarrastes,  which  is  a 
common  species  in  Sumatra.  No  doubt  it  was  on  this  account  that  Fruhstorfer  stated  that  the  species  was 
taken  in  great  numbers  by  Dr  Martin.  It  is  not  possible  to  say  whether  the  figure  of  a  female  in  Seitz 
represents  battakanus  or  sarrastes. 

I  designate  as  lectotype  the  male  in  BMNH,  unfortunately  lacking  its  head,  which  is  labelled  /Type 
[red]/Type  [Fruhstorfer  orange]/CMB  II.  95/Fruhstorfer  Coll.  B.M.  1933-131/Allotinus  battakanus  Frhst. 
[in  Fruhstorfer's  hand]/. 

DISTRIBUTION.  Known  only  from  the  Battak  Mts. 

Allotinus  (Allotinus)  major  C.  &  R.  Felder  sp.  rev. 
(Fig.  8,  Cf  genitalia) 

Allotinus  major  C.  &  R.  Felder,  [1865]:  286,  partim  cf  nee  $,  pi.  35,  fig.  29  cf.  LECTOTYPE  cf, 
SULAWESI  (BMNH),  here  designated  [examined]. 


18  J-   N.  ELIOT 

Allotinus  fallax  major  Felder;  Fruhstorfer,  1913:  343;  1916:  809. 

Allotinus  fallax  depictus  Fruhstorfer,  1913:  343;  1916:  809  [misspelled  depista],  pi.  141h  cf  [as  major]. 

LECTOTYPE  cf ,  SULAWESI  (BMNH),  here  designated  [examined].  Syn.  n. 
Allotinus  kalawarus  Ribbe,  1926:  91.  Syntypes,  SULAWESI  (probably  in  SMT). 

The  most  remarkable  feature  of  this  species  and  A.  maximus  is  the  short  abdomen,  which  in  the  male  does 
not  protrude  beyond  the  hindwings;  this  is  due  to  all  the  abdominal  segments  being  relatively  short 
compared  with  those  of  the  other  species  of  the  genus.  Both  species  can  also  be  recognised  by  the 
arrangement  of  the  postdiscal  series  on  the  underside  of  the  hindwing,  wherein  the  spot  in  space  6  is 
midway  between  those  in  spaces  7  and  5. 

A.  major  shows  much  individual  variation  in  size  and  pattern.  The  male  is  typically  of  medium  size 
(forewing  17-0  mm)  with  a  small,  sullied,  white  patch  on  the  forewing  astride  the  base  of  vein  Cu2. 
Occasionally  the  white  area  below  vein  Cu2  is  unsullied  and  up  to  3-0  mm  wide;  but  in  more  than  half  of  the 
examples  examined  the  white  patch  is  entirely  absent,  and  such  examples  from  east  and  south  Sulawesi 
were  treated  as  a  distinct  subspecies,  depictus,  by  Fruhstorfer.  However,  similar  males  occur  throughout 
Sulawesi,  so  that  depictus  can  only  be  used  as  a  varietal  name.  Females  have,  on  average,  a  larger  white 
patch  than  males.  In  the  commonest  form  the  ovate  white  patch  is  about  4-0  mm  wide  and  fills  the  centre  of 
space  Ib,  the  basal  half  of  space  2  and  may  enter  space  3.  Unmarked  brown  females  appear  to  be  very  rare, 
as  there  is  only  one  example  in  BMNH,  from  west  central  Sulawesi,  without  a  trace  of  white.  Other  females 
from  the  same  area  and  also  from  east  Sulawesi  form  a  complete  connecting  series  leading  up  to  the  normal 
white-patched  form. 

I  have  not  seen  Ribbe's  type-series  of  A.  kalawarus,  so  cannot  be  positive  that  the  name  is  applicable  to 
A.  major.  Ribbe  described  his  species  from  at  least  four  examples  which  he  did  not  sex.  He  wrote  that  the 
upperside  agreed  with  the  figure  of  major  (Fruhstorfer,  1916:  pi.  141h)  which  is  cf-var  depictus  and  that  the 
underside  was  nearest  to  the  figure  of  pyxus  [A.  portunus pyxus]  (Fruhstorfer,  1916:  pi.  141i),  but  differed 
in  having  a  light  whitish  ground  colour  and  heavier  marginal  flecks  and  other  markings.  He  queried 
whether  kalawarus  might  not  be  the  same  as  damodar  (=  A.  macassarensis  macassarensis),  but  said  that  the 
only  specimen  which  he  had  as  damodar  was  a  female  with  a  crenulate  hindwing  termen  (which  is  indeed  a 
character  of  macassarensis  female),  whilst  in  kalawarus  the  termen  was  not  crenulate.  A  non-crenulate 
hindwing  is  a  feature  of  both  sexes  of  A .  major,  so  that  if  any  of  Ribbe's  specimens  were  females  this  would 
confirm  that  they  were  major.  If  they  were  males  the  absence  of  any  mention  of  a  brand  (conspicuous  in  A . 
macassarensis)  also  suggests  that  they  were  major.  I  therefore  feel  reasonably  confident  that  kalawarus  is  a 
synonym  of  major. 


Fig.  8    Allotinus  (Allotinus)  major  C.  &  R.  Felder;  south-west  Sulawesi.  Male  genitalia.  Lower  left, 
uncus,  tegumen  and  brachium  of  an  example  from  north-east  Sulawesi:  Minahassa. 


THE  MILETINI  19 

In  the  male  genitalia  the  uncus/tegumen  blades  are  shorter  and  more  rounded  than  in  any  other  Oriental 
species  of  Miletini.  Judging  by  the  few  preparations  I  have  made  they  are  roundest  in  south  Sulawesi  and 
most  elongated  in  Minahassa,  those  from  east-central  Sulawesi  being  intermediate. 

I  designate  as  lectotype  of  major  a  male  in  BMNH  labelled  /Celeb  Lorquin  type  [blue  circular]/Allotinus 
major  cf  Fd./Type  [red]/FELDER  COLLN. /Rothschild  Bequest  1939-1/.  The  female  figured  by  the 
Felders  is  an  example  of  the  next  species. 

I  designate  as  lectotype  of  depictus  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Sud-Celebes  Dr.  Martin  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/major  fa  depicta  Fr.  [in  Fruhstor- 
fer's  hand]/.  A  female  labelled/Type  [red]/Type  [Fruhstorfer  orange]/O.  Celeb.  Fruhstorfer/depictus  Fruh. 
[in  Corbet's  hand]/  is  a  paralectotype;  it  is  a  typical  example  with  a  white  patch  on  the  forewing. 

DISTRIBUTION.  The  species  occurs  at  low  levels  throughout  Sulawesi  and  in  Banggai  I.  and  Sangihe  I. 

Allotinus  (Allotinus)  maximus  Staudinger  stat.  n. 
(Fig.  9,  cf  genitalia) 

[Allotinus  major  C.  &  R.  Felder,  [1865]:  286,  partim  $  nee  cf ,  pi.  35,  figs  30,  31  $.  Misidentification.] 
Allotinus  albatus  Felder  var.  maximus  Staudinger,  1888:  269,  pi.  94.  Holotype  cf,  SULAWESI  (coll. 

Staudinger,  probably  in  MNHU). 
Allotinus  fallax  major  Felder  $-f.  albatus  Felder  [=  maximus};  Fruhstorfer,  1913:  343;  1916:  809 

[misspelled  albadus  and  magnimus}. 

Both  Fruhstorfer  and  Staudinger  confused  this  species  with  Allotinus  albatus,  a  distinct  species  in  subgenus 
Paragerydus  (p.  30).  The  former  recognised  its  kinship  with  A.  major,  regarding  it  as  a  montane  variety 
from  Minahassa.  I  think  it  is  more  likely  that  major  and  maximus  are  distinct  sibling  species.  The  latter  may 
well  be  wholly  montane,  since  of  four  males  in  BMNH  the  only  two  with  complete  data  came  from  the  Peak 
of  Bonthain,  1000-2000  m.  These  examples  from  the  extreme  south-west  of  Sulawesi  do  not  appear  to 
differ  from  Staudinger's  holotype  male  from  Minahassa,  so  that  it  is  probable  that  the  species  occurs 
unchanged  throughout  the  mountains  of  Sulawesi. 

A.  maximus  is  larger  than  A.  major,  and  in  the  male  there  is  a  clear  white  patch  on  the  forewing  filling  the 
basal  half  of  space  2  and  the  central  half  of  space  1  b ,  which  is  grey  at  the  wing  base .  The  only  female  I  have 
seen  is  the  misidentified  female  syntype  of  major  figured  by  the  Felders,  which  has  a  large  white  patch  on 
the  forewing  extending  into  the  base  of  space  4  and  the  lower  edge  of  the  cell.  It  is  labelled  /Type 
[red]/Celebes  Lorquin  type  [blue  circular]/Allotinus  major  n  $  Fd. /major  n/FELDER  COLLN. /Roth- 
schild Bequest  1939-1/. 

The  male  genitalia  (Fig.  9)  of  the  single  example  dissected  differ  slightly  from  those  of  A.  major  in  having 
a  narrower,  more  pointed  phallus  and  longer,  more  rectangular  uncus  blades. 

DISTRIBUTION.  Confined  to  Sulawesi. 


Fig.  9    Allotinus  (Allotinus)  maximus  Staudinger;  Sulawesi  (no  further  data).  Male  genitalia. 


20  J.   N.  ELIOT 

FABITARAS  subgen.  n. 

Type-species:  Paragerydus  fabius  Distant  &  Pryer,  1887:  266.  Gender  masculine. 

Antenna  nearly  two-thirds  length  of  forewing  costa,  with  about  62-66  segments.  In  males,  abdominal  hair 
tufts  short,  not  protruding  except  when  genitalia  are  everted.  Forewing  venation  particularly  subject  to 
individual  variation,  with  apical  angle  of  cell  sometimes  sharply  acute,  sometimes  almost  a  right  angle. 
Veins  Ml  and  R5  usually  stalked,  but  sometimes  connate  from  cell  apex,  most  often  in  A.  taras  and  A. 
strigatus,  while  in  A.  bldiensis  they  may  be  just  separate.  Vein  RT,  also  very  variable,  usually  arising  from 
about  middle  of  vein  R5,  but  sometimes  as  short  as  in  subgen.  Paragerydus,  sometimes  as  long  as  in  typical 
species  of  subgen.  Allotinus.  In  males  vein  M3  always  basally  swollen  and  clothed  with  small  specialised 
scales.  Hindwing  usually  with  a  weakly  developed  humeral  vein  extending  less  than  half-way  across  space 
8,  but  sometimes  absent  or  vestigial,  most  often  in  females.  Underside  characterised  by  a  series  of 
submarginal  blackish  dots  which,  on  the  forewing,  are  outwardly  white-edged  in  females  but  not,  or  more 
indistinctly  so,  in  males;  indeed,  even  the  blackish  dots  may  be  absent  in  males  of  A.  strigatus  and  A. 
portunus.  Male  genitalia  not  unlike  those  of  subgen.  Allotinus,  with  valva  ending  in  an  apical  hook  but 
edges  folded  inwards  more  strongly  in  the  form  of  a  trough. 

The  species  included  in  Fabitaras  were  placed  in  Paragerydus  by  earlier  authors  up  to  and  including 
Fruhstorfer  (1916)  because  of  the  swollen  vein  M3  of  males  and  the  common  origin  of  veins  M}  and  R5  in 
both  sexes;  but  on  the  basis  of  the  male  genitalia  and  the  frequent  presence  of  a  humeral  vein  it  is  certain 
that  Fabitaras  is  more  closely  allied  to  Allotinus  than  to  Paragerydus. 

The  subgenus  is  found  from  central  Burma  to  Sundaland  and  the  Philippines,  and  comprises  ten  species. 

Key  to  the  species  of  subgenus  Fabitaras 

1  Underside  of  hindwing  with  postdiscal  spot  in  space  6  placed  below,  or  just  inside,  that  in  space 

7.  Female  with  hindwing  produced  at  vein  M3  (faWus-group) 2 

Underside  of  hindwing  with  postdiscal  spot  in  space  6  placed  well  outside  spot  in  space  7.  Female 

hindwing  rounded  (faras-group) 4 

2  Underside  of  forewing  with  postdiscal  series  more  or  less  parallel  to  termen  except  that  spot  in 

space  5  is  moved  a  little  outwards.  Postdiscal  spots  above  vein  M3  on  forewing  and  vein  M\  on 

hindwing  not  enlarged  nor  blotchy 3 

Underside  of  forewing  with  postdiscal  series  inclined  towards  apex;  spots  in  spaces  4  and  5 
moved  out  of  line,  the  latter  very  close  to  termen.  Postdiscal  spots  above  vein  M3  on  forewing 
and  vein  MI  on  hindwing  enlarged  and  blotchy fabius  (p.  21) 

3  cf  upperside  of  forewing  with  vein  M3  swollen  for  three-quarters  of  its  length;  visual  brand 

prominent.  Underside  of  forewing  with  apical  region  shaded  reddish  brown;  hindwing  with 
submarginal  black  spot  in  space  6  at  most  barely  larger  than  other  submarginal  spots 

borneensis  (p.  22) 

Cf  upperside  of  forewing  with  vein  M3  swollen  for  half  its  length;  visual  brand  narrow  and 
obscure.  Underside  of  forewing  with  apical  region  not  shaded  reddish  brown;  hindwing  with 
submarginal  black  spot  in  space  6  triangular  and  larger  than  other  submarginal  spots 

punctatus  (p.  23) 

4  Cf  9  upperside  of  hindwing  with  tornal  area  brown 5 

Cf  9  upperside  of  hindwing  with  tornal  quarter  white brooksi  (p.  26) 

5  Underside  of  hindwing  with  submarginal  black  spot  in  space  6  at  most  barely  larger  than  other 

submarginal  spots,  cf  upperside  of  forewing  with  at  least  indications  of  a  visual  brand  astride 

swollen  portion  of  vein  M3 6 

Underside  of  hindwing  with  submarginal  black  spot  in  space  6  triangular  and  larger  than  other 
submarginal  spots,  cf  upperside  of  forewing  without  a  visual  brand.  9  upperside  of  hindwing 
may  have  a  brownish  cream  border nigritus  (p.  24) 

6  Underside  of  hindwing  with  postdiscal  spot  in  space  6  placed  roughly  midway  between  the  spots 

in-spaces  7  and  5 7 

Underside  of  hindwing  with  postdiscal  spot  in  space  6  placed  much  further  from  spot  in  space  7 
so  that  it  is  almost  on  an  even  arc  with  the  spots  in  spaces  2  to  5 8 

7  Underside  ground  colour  white,  cf  forewing  with  vein  M3  swollen  for  only  one-third  its  length; 

visual  brand  ill-defined  and  less  than  1  -0  mm  wide bidiensis  (p.  26) 

Underside  ground  colour  pale  buff,  cf  forewing  with  swollen  portion  of  vein  M3  about  half  its 

length;  visual  brand  about  2-0  mm  wide strigatus  (p.  24) 

8  cf  forewing  with  vein  M3  swollen  for  more  than  half  its  length ;  visual  brand  at  least  1-0  mm  wide  9 


THE  MILETINI  21 

O"  forewing  with  vein  M3  swollen  for  half  its  length;  visual  brand  very  narrow,  inconspicuous. 

Underside  ground  colour  greyish  white  ;forewing  apex  shaded  reddish  brown turns  (p.  27) 

9  Underside  ground  colour  greyish  white ,  not  darkened  towards  forewing  apex;  black  submargin- 
al  spots  outwardly  white-edged;  postdiscal  series  slightly  closer  to  termen  at  dorsum  than  at 
costa.  cf  upperside  of  forewing  with  vein  M3  swollen  for  two-thirds  its  length;  visual  brand  up 

to  2-0  mm  wide,  comparatively  well  defined sarrastes  (p.  28) 

-  Underside  ground  colour  in  cf  buff,  becoming  darker  towards  forewing  apex;  striation  dense, 
black  marginal  spots  absent  or  obscure,  not  white-edged.  $  ground  colour  greyish  white  with 
forewing  apex  more  or  less  shaded  with  brown;  submarginal  white-edged  black  spots  present 
on  forewing.  Postdiscal  series  parallel  to  termen.  cf  upperside  of  forewing  with  vein  M3 
swollen  for  nearly  four-fifths  its  length;  visual  brand  ill-defined,  just  over  1-0  mm  wide 

portunus  (p.  29) 

Allotinus  (Fabitaras)  fabius  (Distant  &  Pryer) 

(Fig.  10,  cf  genitalia) 
Paragerydus  fabius  Distant  &  Pryer,  1887:  266. 

The  species  is  readily  recognised  by  the  underside  markings.  On  the  forewing  the  postdiscal  spots  in  spaces 
4  and  especially  5  are  shifted  towards  the  termen,  whilst  the  spots  in  spaces  6  and  7  are  placed  much  further 
basad;  in  addition  the  spots  above  vein  M3  are  more  or  less  enlarged  and  blotchy.  On  the  hindwing  the 
postdiscal  spots  in  spaces  6  and  7  are  placed  one  above  the  other  and  are  large  and  blotchy.  The  female  has 
the  hindwing  weakly  caudate  at  vein  M3.  The  male  has  vein  M3  of  the  forewing  swollen  for  about 
three-quarters  of  its  length  and  the  visual  brand  is  about  1-5  mm  wide. 

The  species  has  a  restricted  distribution  in  Sundaland,  and  has  not  been  found  in  Java,  Palawan  nor  the 
islands  off  the  west  coast  of  Sumatra.  There  are  two  subspecies. 

Key  to  the  subspecies  of  A.  (F.)  fabius 

1     $  upperside  of  hindwing  outwardly  white fabius  fabius  (p.  21) 

$  upperside  of  hindwing  brown fabius  arrius  (p.  21) 

Allotinus  (Fabitaras)  fabius  fabius  (Distant  &  Pryer) 

Paragerydus  fabius  Distant  &  Pryer,  1887:  266;  Cowan,  1966: 5.  Holotype  $ ,  BORNEO:  Sandakan  (BMNH) 

[examined]. 
Allotinus  caudatus  Grose-Smith,  1893:  34.  Holotype  'cf'  recte  $,  BORNEO:  Mt  Kina  Balu  (BMNH) 

[examined].  Syn.  n. 

Paragerydus  caudatus  (Grose-Smith);  H.  H.  Druce,  1895:  563,  pi.  31,  figs  7,  8  cf  - 
Allotinus  fabius  fabius  (Distant  &  Pryer);  Fruhstorfer,  1914:  22;  1916:  814;  Corbet,  19396:  74. 
Allotinus  fabius  caudatus  Smith;  Fruhstorfer,  1914:  22;  1916:  814;  Corbet,  19396:  74. 
Allotinus  fabius  pamisus  Fruhstorfer,  1914:  22;  1916:  814;  Corbet,  19396:  74  [misspelt  pamiscus].  Holotype 

$,  BORNEO:  south-east  (BMNH)  [examined].  Syn.  n. 

Fruhstorfer  (1915)  perpetuated  Grose-Smith's  mistake  over  the  sex  of  caudatus  by  incorrectly  stating  that 
both  sexes  of  the  race  from  Kina  Balu  have  the  outer  part  of  the  hindwing  white.  In  fact  it  is  only  females 
which  have  the  hindwing  partly  white,  and  examples  from  Kina  Balu  do  not  differ  significantly  from  those 
from  the  rest  of  Borneo  nor  from  examples  from  south-west  Sumatra. 

DISTRIBUTION.  Borneo,  including  Pulo  Laut;  south  Sumatra  (1  cf ,  3  $ ,  Lebong  Tandai  (C.  J.  Brooks)). 

Allotinus  (Fabitaras)  fabius  arrius  Fruhstorfer 
(Fig.  10,  cf  genitalia) 

[Paragerydus panormis  Elwes,  1893:  619  (partim),  pi.  43,  fig.  9  $.  Misidentification.] 

Allotinus  fabius  arrius  Fruhstorfer,  1914:  22;  1916:  814,  pi.  1411,  cf  $;  Corbet,  19396:  74,  fig.  4  cf  genitalia; 

Fleming,  1975:  21,  pi.  58,  fig.  L45  cf;  Eliot,  1978:  240.  LECTOTYPE  cf.  SUMATRA  (BMNH),  here 

designated  [examined]. 
[Allotinus  fabius  panormis  (Elwes);  Fruhstorfer,  1916:  814.  Misidentification.] 


22 


J.  N.  ELIOT 


Fig.  10    Allotinus  (Fabitaras)  fabius  arrius  (Fruhstorfer);  Malay  Peninsula.  Male  genitalia. 

Differs  from  the  nominate  subspecies  only  in  the  all  brown  female. 

I  designate  as  lectotype  of  arrius  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/CMB 
II.  94/Sumatra  Monies  Batlak  ex  coll.  Fruhslorfer/Fruhstorfer  Coll.  B.M.  1933-131/fabius  arrius  Frhsl.  [in 
Fruhslorfer's  hand]/.  A  similarly  labelled  female  is  a  paraleclolype. 

DISTRIBUTION.  Malay  Peninsula;  north  Sumatra. 

Allotinus  (Fabitaras)  borneensis  Moulton 
(Figs  11  Cf  genitalia;  58  cf) 

Allotinus  borneensis  Moullon,  1911:  81;  Fruhslorfer,  1913:  371;  1916: 814.  Hololype  cf ,  BORNEO:  Sarawak 

(BMNH)  [examined]. 

Allotinus  borneensis  borneensis  Moullon;  Corbel,  19396:  76,  fig.  3,  cf  genitalia. 
Allotinus  borneensis  elioti  Corbet,  19396:  76;  Fleming,  1975:  22,  pi.  58,  fig.  L49  cf ;  Eliol,  1978:  240. 

Hololype  cf ,  WEST  MALAYSIA  (BMNH)  [examined].  Syn.  n. 

On  Ihe  upperside  bolh  sexes  resemble  A.  fabius  arrius,  but  are  more  reddish  brown  and,  in  Ihe  male,  Ihe 


Fig.  11     Allotinus  (Fabitaras)  borneensis  Moulton;  Malay  Peninsula.  Male  genitalia. 


THE  MILETINI 


23 


swollen  portion  of  vein  M3  is  very  slightly  longer  and  the  visual  brand  is  very  slightly  wider.  The  underside  is 
greyish  white  to  pale  buff-white,  with  the  apical  area  of  the  forewing  shaded  with  reddish  brown.  On  the 
forewing  the  postdiscal  series  is  parallel  to  the  termen  except  that  the  spot  in  space  5  is  shifted  outwards,  but 
not  nearly  as  much  as  in  A.  fabius.  On  the  hindwing  the  spot  in  space  6  is  usually  placed  just  inside  that  in 
space  7. 

DISTRIBUTION.  Borneo;  Malay  Peninsula;  Sumatra;  Bangka. 

Allotinus  (Fabitaras)  punctatus  (Semper) 
(Figs  12,  cf  genitalia;  59  9) 

Paragerydus  punctatus  Semper,  1889:  165,  pi.  31,  figs  16  cf,  17  $.  LECTOTYPE  cf,  PHILIPPINES: 
Mindanao  (SM),  here  designated  [examined]. 

Allotinus  punctatus  (Semper);  Fruhstorfer,  1913:  371;  1916:  814;  Corbet,  1939b:  74,  fig.  1,  cf  genitalia. 

Allotinus  anaxandridas  Fruhstorfer,  1916:  814.  Holotype  9,  PHILIPPINES:  Mindanao  (SM)  [examined]. 
[$-morph.]  Syn.  n. 

Allotinus  caesemius  Fruhstorfer,  1916:  814.  LECTOTYPE  $,  PHILIPPINES:  Mindanao  (SM),  here  desig- 
nated [examined].  [$-morph.]  Syn.  n. 

On  the  upperside  the  male  has  vein  M3  swollen  for  half  its  length  and  the  visual  brand  is  rather  narrow  and 
ill-defined.  The  underside  is  pale  buff  with  the  postdiscal  markings  showing  great  individual  variation  in 
width,  in  some  examples  being  as  much  as  2-0  mm  wide.  In  females  the  ground  colour  is  greyish  white,  and 
the  postdiscal  markings  are  much  narrower,  seldom  as  much  as  1-0  mm  in  width.  The  white-edged  black 
submarginal  spots  are  well  marked,  that  in  space  6  of  the  hindwing  being  triangular  and  larger  than  the 
others.  As  in  A.  fabius  and  A.  borneensis  the  postdiscal  spots  in  spaces  7  and  6  of  the  hindwing  are  placed 
more  or  less  one  above  the  other,  and  the  female  has  the  hindwing  weakly  caudate  at  vein  M3. 

The  species  is  chiefly  remarkable  for  the  polymorphism  of  the  female,  which  was  recognised  by  Semper 
who  described  three  morphs.  Fruhstorfer,  however,  considered  that  the  three  morphs  were  distinct  species 
-  a  view  for  which  there  is  no  supporting  evidence.  The  typical  female  is  plain  brown  on  both  wings.  In  $-f. 
caesemius  Fruhstorfer  the  forewing  has  a  large  white  patch  while  the  hindwing  is  brown;  it  seems  to  be  the 
commonest  morph.  In  $-f.  anaxandridas  Fruhstorfer  the  forewing  resembles  that  of  caesemius  but  the 
hindwing  also  has  a  large  white  patch;  it  appears  to  be  much  the  rarest  morph.  Another  9 -morph  was 
described  by  Fruhstorfer  as  9-f-  eretria,  but  I  think  that  the  butterfly  in  question  was  a  female  of  A.  nigritus 
(seep.  24). 

Semper  did  not  designate  a  type  of  punctatus,  whilst  Fruhstorfer  named  caesemius  from  two  females  and 
anaxandridas  from  a  single  female  in  Semper  coll.,  which  Semper  had  designated  merely  as  varieties  of 
punctatus.  I  now  designate  as  lectotype  of  punctatus  one  of  two  males  ex  Semper  coll.  in  SM  labelled  /Coll. 
C.  Semper/Sibulan/215/Parag.  punctatus  typ.  Semper/297c/Typus  [red]/.  A  second  male  labelled  /215/Coll. 
C.  Semper/  is  a  paralectotype.  I  designate  as  lectotype  of  caesemius  a  female  ex  Semper  coll.  in  SM  labelled 
/Sibulan/297a/215/.  The  single  female  of  anaxandridas  is  automatically  the  holotype  and  has  been  so 
labelled. 


Fig.  12    Allotinus  (Fabitaras)  punctatus  (Semper);  Mindanao.  Male  genitalia. 


24  J.  N.  ELIOT 

DISTRIBUTION.  The  species  has  hitherto  been  recorded  only  from  Mindanao.  There  is  a  single  female  in  coll. 
Treadaway  from  Leyte  (Catmon  450  m,  10. v.  1977)  which  differs  from  f.  caesemius  in  having  the  white 
forewing  patch  reduced,  with  the  dusky  scaling  at  the  wing  base  and  above  the  dorsum  almost  as  dark 
brown  as  the  marginal  and  costal  borders,  whilst  the  underside  is  yellowish  white  with  the  markings  heavier 
and  more  reddish  than  in  females  from  Mindanao.  It  is  probable  that  a  distinct  subspecies  flies  in  Leyte,  but 
I  hesitate  to  name  one  on  the  basis  of  a  single  female. 

Allotinus  (Fabitaras)  nigritus  (Semper) 
(Figs  13,  Cf  genitalia,  60  $) 

Paragerydus  nigritus  Semper,  1889: 164,  pi.  31,  fig.  15  cf .  LECTOTYPE  cf ,  PHILIPPINES:  Mindanao  (SM), 

here  designated  [examined]. 

Allotinus  nigritus  (Semper);  Fruhstorfer,  1914:  22;  1916:  814. 
Allotinus punctatus  (Semper)  $-f.  eretria  Fruhstorfer,  1916:  814.  Holotype  $.  PHILIPPINES:  Mindanao  (not 

located). 

In  the  male  vein  M3  is  swollen  for  a  little  under  half  its  length  and  there  is  no  visual  brand.  The  underside  is 
pale  brownish  ochreous  with  darker  brown  markings.  The  black  submarginal  spots  resemble  those  of  A. 
punctatus,  from  which  A.  nigritus  can  be  separated  by  the  postdiscal  spot  in  space  6  of  the  hindwing  being 
placed  midway  between  the  spots  in  spaces  7  and  5,  or  a  little  closer  to  the  latter,  as  well  as  by  its  richer, 
more  ochreous  appearance. 

The  only  female  I  have  seen  (Mindanao,  Mt  Apo,  15. ii.  1983,  (A.  Ballantine))  is  dark  brown  above  with 
an  inwardly  diffuse,  sullied  whitish  border  2-0  mm  wide  on  the  hindwing  from  the  tornus  to  vein  R5.  The 
hindwing  termen  is  barely  dentate  at  the  end  of  vein  M3,  as  in  all  species  of  the  subgenus  dealt  with 
subsequently. 

Fruhstorfer  named  as  A.  punctatus  $  -f .  eretria  a  female  with  the  underside  bright  ochreous  bearing  thick 
brown  markings  and  black  marginal  spots.  The  underside  of  females  of  A.  punctatus  is  greyish  white,  so 
that  I  suspect  that  eretria  really  applies  to  A  nigritus.  Fruhstorfer  did  not  mention  the  upperside  of  eretria, 
which  was  presumably  unmarked  brown,  as  in  the  typical  female  morph  of  A.  punctatus,  and  if  my 
supposition  of  the  true  identity  of  eretria  is  correct  it  would  appear  that  the  female  of  A.  nigritus  is 
dimorphic  -  with  or  without  a  whitish  border  on  the  hindwing. 

I  designate  as  lectotype  of  nigritus  a  male  in  SM  labelled  /Coll.  C.Semper/Ost  Mind./214/297b/15/Parag. 
nigritus  typ.  Semper/Typus  [red]/.  A  second  male  labelled  /Ost  Mind./214/Typus  [red]/is  a  paralectotype. 

DISTRIBUTION.  Mindanao. 


Fig.  13    Allotinus  (Fabitaras)  nigritus  (Semper);  Mindanao.  Male  genitalia. 

Allotinus  (Fabitaras)  strigatus  Moulton 

(Figs  14,  cf  genitalia;  61  cf ) 
Allotinus  strigatus  Moulton,  1911:  80. 
The  species  can  be  recognised  by  the  fact  that  on  the  underside  of  the  hindwing  the  postdiscal  spot  in  space 


THE  MILETINI 


25 


6  is  placed  midway  between  those  in  spaces  7  and  5,  while  on  the  forewing  the  postdiscal  series  is  much 
closer  to  the  termen  near  the  tornus  than  near  the  costa.  The  underside  ground  colour  is  a  uniform  pale 
buff.  On  the  upperside  the  male  has  vein  A/3  swollen  for  half  its  length,  and  the  visual  brand  is  quadrate, 
about  2  mm  wide  and  not  sharply  outlined. 

The  species  is  strictly  Sundanian,  and  has  not  yet  been  found  in  Java,  Palawan  or  the  islands  off  the  west 
coast  of  Sumatra.  There  are  two  subspecies. 

Key  to  the  subspecies  of  A.  (F.)  strigatus 

1  On  the  underside  of  the  forewing  the  blackish  submarginal  dots  are  inconspicuous  and  not 

outwardly  white-edged strigatus  strigaius  (p.  25) 

-  On  the  underside  of  the  forewing  the  blackish  submarginal  dots  are  outwardly  white-edged  in  $ 

and  in  the  apical  half  of  the  wing  in  cf strigatus  malayanus  (p.  25) 


Allotinus  (Fabitaras)  strigatus  strigatus  Moulton 
(Fig.  61  cf) 

Allotinus  strigatus  Moulton,  1911:  80.  Holotype  cf ,  BORNEO:  Pulo  Laut  (BMNH)  [examined]. 
Allotinus  strigatus  strigatus  Moulton;  Fruhstorfer,  1914:  22;  1916:  813;  Corbet,  1939ft:  75,  fig.  5,  cf 
genitalia. 

On  the  underside  the  submarginal  blackish  dots  are  not  white-edged,  and  the  usual  markings  are 
comparatively  broad  and  well  defined. 

DISTRIBUTION.  Borneo,  including  Pulo  Laut. 


Allotinus  (Fabitaras)  strigatus  malayanus  Corbet 
(Fig.  14,  cf  genitalia) 

Allotinus  strigatus  malayanus  Corbet,  1939ft:  75;  Fleming,  1975:  22,  pi.  58,  fig.  L48  cf ;  Eliot,  1978:  240. 

Holotype  cf ,  WEST  MALAYSIA  (BMNH)  [examined]. 
Allotinus  strigatus  denalus  Corbet,  1939ft:  75.  Holotype  cf ,  SUMATRA:  Battak  Mts  (BMNH)  [examined]. 

Syn.  n. 

On  the  underside  of  the  forewing  the  blackish  submarginal  dots  are  outwardly  edged  with  whitish  in  the 
female,  but  in  the  male  less  distinctly  and  only  in  the  apical  half  of  the  wing.  The  markings  are  narrower  and 
less  well  defined  than  in  the  nominate  subspecies.  In  the  male  the  swelling  of  vein  M3  is  slightly  shorter, 
being  just  under  half  its  length,  and  the  visual  brand  is  usually  a  little  narrower. 

DISTRIBUTION.  Malay  Peninsula,  including  Singapore;  Sumatra. 


Fig.  14    Allotinus  (Fabitaras)  strigatus  malayanus  Corbet;  Malay  Peninsula.  Male  genitalia. 


26 


J.  N.  ELIOT 


Allotinus  (Fabitaras)  brooksisp.  n. 

(Figs  15,  cf  genitalia;  63  cf ) 

Cf  forewing  length  15-0  mm.  Venation  normal  for  the  subgenus,  with  veins  M j  and  R5  of  forewing  having  a 
moderately  long  common  stalk.  Upperside  brown,  with  tornal  quarter  of  hindwing  white,  very  sparsely 
dusted  with  brown  scales  and  with  adjoining  cilia  white  chequered  with  dark  brown  at  vein  endings. 
Forewing  with  vein  M3  swollen  for  slightly  less  than  one-third  of  its  length;  visual  brand  very  narrow, 
obscure.  Underside  very  pale  greyish  white,  sparsely  striated;  postdiscal  spots  small ,  more  or  less  parallel 
to  termen;  hindwing  with  postdiscal  spot  in  space  6  a  little  closer  to  that  in  space  7  than  to  that  in  space  5; 
cell-end  bars  unusually  heavy,  especially  on  hindwing;  forewing  with  submarginal  blackish  dots  outwardly 
white-edged.  Genitalia  similar  to  those  of  A.  strigatus,  A.  taras  and  A.  sarrastes,  but  apical  process  of  valva 
slightly  larger. 

$  forewing  length  15-0  mm.  Differs  from  male  in  that  inwardly  diffuse  white  area  on  hindwing  a  little 
larger,  occupying  one-third  of  wing.  At  first  sight  this  sex  might  be  mistaken  for  A.  fabius  fabius ,  but  the 
rounded  hindwing  termen  separates  it  readily. 

MATERIAL  EXAMINED 

Holotype  cf ,  Borneo:  'Bau  Feb  Feb  10',  'Sarawak  C.  J.  Brooks',  '192',  'C.  J.  Brooks  Bequest.  B.M. 
1953-173'  (BMNH). 

Paratype.  Borneo:  1  $  (allotype),  'Bau  Dec.  09',  '192',  'C.  J.  Brooks  Bequest  B.M.  1953-173'  (BMNH). 


Fig.  15    Allotinus  (Fabitaras)  brooksi  sp.  n. ;  Borneo.  Male  genitalia. 


Allotinus  (Fabitaras)  bidiensissp.  n. 

(Figs  16,  cf  genitalia;  62  cf ) 

Cf  forewing  length  15-0-16-0  mm.  In  the  two  examples  examined  forewing  venation  differs  from  rest  of 
subgenus  in  that  veins  MI  and  R5  are  just  separate  at  their  origins;  vein  /?3  rather  short,  arising  just  before 
end  of  vein  R2.  Upperside  brown,  with  basal  third  of  vein  M3  swollen;  visual  brand  narrow,  obscure. 
Underside  with  markings  generally  arranged  as  in  sympatric  A.  strigatus,  but  differing  in  the  pale  greyish 
white  ground  colour,  the  smaller,  more  rounded  postdiscal  spots  and  the  prominently  white-edged  blackish 
submarginal  dots  on  the  forewing.  Male  genitalia  distinguished  by  valva,  in  which  the  lower  process 
extends  beyond  the  short  apical  hook,  and  comparatively  short,  stout  phallus. 

9  forewing  length  14-0  mm  in  the  single  example  seen,  wherein  veins  MI  and  R5  of  forewing  connate 
from  cell  apex.  Upperside  brown.  Underside  similar  to  male. 

MATERIAL  EXAMINED 

Holotype  cf ,  Borneo:  Sarawak,  '647  20/5/08',  '190f ,  'C.  J.  Brooks  Bequest.  B.M.  1953-173'  (BMNH). 

Paratypes.  Borneo:  1  $  (allotype),  Sabah,  'N.  Born.',  'Madai  3.2.92',  'Joicey  Bequest.  Brit.  Mus. 
1934-120'  (BMNH);  cf ,  'Bidi  Sarawak  1907  C.  J.  Brooks',  '190f ,  'C.  J.  Brooks  Bequest.  B.M.  1953-173' 
(BMNH). 


THE  MILETINI 


27 


Fig.  16    Allotinus  (Fabitaras)  bidiensis  sp.  n. ;  Borneo.  Male  genitalia. 


Allotinus  (Fabitaras)  taras  (Doherty) 

(Fig.  17,  Cf  genitalia) 

Paragerydus  taras  Doherty,  1889:  437,  pi.  23,  fig.  10  cf .  Syntypes,  BURMA:  Tenasserim  (not  located). 
Parageryduspanormis  Elwes,  1893:  619,  partim  'cf '  recte  £ ,  nee  $ ,  pi.  43,  fig  8  'cf '.  Holotype  'cf '  recte  $ , 

BURMA:  East  Pegu  (BMNH)  [examined].  Syn.  n. 
Allotinus  taras  (Doherty)  Bingham,  1907:  300;  Swinhoe,  1910:  199,  pi.  617,  figs  2,  2b  cf ,  2a,  2c  $;  Evans, 

1932:  212;  Cantlie,  1963:  27. 
Allotinus panormis  (Elwes)  Bingham,  1907:  301;  Swinhoe,  1910: 197  partim,  pi.  616,  figs 3, 3b  'cf'  recte  $, 

nee  $. 

Allotinus  taras  taras  (Doherty);  Fruhstorfer,  1913:  370;  1916:  813;  Corbet,  19396:  74,  partim. 
Allotinus  fabius panormis  (Elwes);  Fruhstorfer,  1916:  814,  partim;  Evans,  1932:  212. 
Allotinus  panormis  panormis  (Elwes);  Corbet,  19396:  74;  Cantlie,  1963:  27. 

On  the  upper  surface  of  the  forewing  the  male  has  vein  M3  swollen  for  half  its  length,  and  the  visual  brand  is 
very  narrow  and  inconspicuous;  indeed  Doherty,  in  his  original  description,  gave  absence  of  the  brand  as 


Fig.  17    Allotinus  (Fabitaras)  taras  (Doherty);  Burma.  Male  genitalia. 


28 


J.   N.   ELIOT 


one  of  the  characters  of  the  species.  The  underside  of  both  sexes  is  greyish  white  turning  to  reddish  brown 
towards  the  forewing  apex  and,  occasionally,  at  the  hindwing  apex  also.  The  postdiscal  series  of  spots  is 
rather  lightly  marked,  and  though  these  spots  were  mentioned  by  Doherty  they  are  wanting  in  the  example 
chosen  for  his  figure.  Such  examples  cannot  be  regarded  as  typical,  only  two  out  of  28  males  in  BMNH 
being  similar  to  Doherty's  figure. 

The  'male'  (recte  female)  of  Paragerydus  panormis  is  a  quite  typical  female  of  A  taras,  and  I  am  at  a  loss 
to  understand  why  Corbet  regarded  it  as  a  different  species  conspecific  with  A.  portunus. 

The  male  genitalia  of  A.  taras  and  A.  sarrastes  are  similar,  so  there  are  grounds  for  regarding  these  two 
taxa  as  conspecific.  However,  they  overlap  in  south  Burma  between  Tavoy  and  Mergui  over  a  distance  of 
some  160  km  without  any  evidence  of  intergradation,  so  that  it  seems  highly  probable  that  their  period  of 
isolation,  when  there  was  a  sea  barrier  in  the  region  of  the  Isthmus  of  Kra,  was  sufficiently  prolonged  for 
interbreeding  to  be  impossible  when  they  again  met. 

I  have  seen  none  of  Doherty's  type-series  and  as  the  figured  male  is  atypical  I  do  not  designate  it  as 
lectotype. 

DISTRIBUTION.  Burma,  from  the  Karen  Hills  to  Mergui. 

Allotinus  (Fabitaras)  sarrastes  Fruhstorfer  stat.  n. 
(Fig.  18,  cf  genitalia) 

Allotinus  taras  sarrastes  Fruhstorfer,  1913:  370;  1916:  813;  Corbet,  19396:  74;  Eliot,  1961:  71;  1978:  240; 

Fleming,  1975:  21,  pi.  58,  fig.  L46  cf .  LECTOTYPE  cf ,  BORNEO  (BMNH),  here  designated  [examined]. 
[Allotinus  taras  battakanus  Fruhstorfer,  1913:  370,  partim;  1916:  813  partim,  ?  pi.  141g  $.  Misidentifica- 

tion.] 

Allotinus porriginosus  Toxopeus,  1932:  Ixxvii.  Holotype  cf ,  JAVA:  south  (not  located).  Syn.  n. 
[Allotinus  taras  taras  (Doherty);  Corbet,  19396:  74,  fig.  2,  cf  genitalia.  Misidentification.] 
Allotinus  taras  mendava  Riley,  1944:  253,  pi.  2,  fig.  30  cf,  31  $.  Holotype  cf,  MENTAWAI  Is.:  Sipora 

(BMNH)  [examined].  Syn.  n. 

This  Sundanian  species  is  obviously  very  closely  related  to  A.  taras,  but  it  is  best  regarded  as  specifically 
distinct.  It  differs  from  A.  taras  on  the  underside  by  the  absence  of  reddish  brown  shading  towards  the 
forewing  apex.  In  addition,  on  the  upperside  of  the  male  forewing  the  swelling  of  vein  M3  extends  over 
about  two-thirds  of  its  length  and  the  visual  brand  is  prominent  and  usually  about  1-5  mm  wide.  On  the 
underside  the  white  edges  to  the  blackish  submarginal  spots  are  particularly  well  marked,  especially  in  the 
female. 

I  designate  as  lectotype  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/Kina  Balu  ex 
coll.  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/sarrastes  Frhst.  [in  Fruhstorfer's  hand]/.  A  female 
labelled  /Type  [red]/Type  [Fruhstorfer  orange]/Nord-Borneo  Brunei  Waterstradt  1890/Fruhstorfer  Coll. 
B.M.  1933-131/sarrastes  Frhst.  [in  Fruhstorfer's  hand]/  is  a  paralectotype. 


Fig.  18    Allotinus  (Fabitaras)  sarrastes  Fruhstorfer;  Malay  Peninsula.  Male  genitalia. 


THE  MILETINI  29 

DISTRIBUTION.  Burma,  from  Tavoy  southwards;  Malay  Peninsula;  Sumatra;  Mentawai  Is.;  Borneo;  Java; 
Mindanao  (SM,  1  cf  labelled  /Ost  Mind./298a/213/Coll.  C.  Semper/re verdini/;  this  specimen  has  the  visual 
brand  obscure  and  ill-defined,  but  this  may  be  due  partly  to  its  rubbed  condition.) 

Allotinus  (Fabitaras)  portunus  (de  Niceville) 

(Fig.  19,  cf  genitalia) 
Paragerydus portunus  de  Niceville,  1894:  27. 

Males  are  easily  recognised  by  the  long  swelling  of  vein  M3  of  the  forewing,  extending  over  nearly 
four-fifths  of  its  length,  and  the  rather  diffuse  and  narrow  visual  brand,  as  well  as  by  the  buff  or  pale  rufous 
underside  on  which  the  usual  blackish  submarginal  spots  are  absent  or,  if  present,  are  not  outwardly 
white-edged.  Females  most  nearly  resemble  that  sex  of  A.  taras  on  the  underside,  but  the  subapical 
darkening  on  the  forewing  is  less  developed  and  brownish  ochreous  without  the  red  tinge  of  taras.  In  both 
sexes  the  postdiscal  series  of  spots  on  the  forewing  is  more  exactly  parallel  to  the  termen  than  in  the  other 
species  of  the  subgenus. 

As  already  pointed  out  (p.  28)  Corbet  (19396)  subordinated  A.  portunus  and  its  subspecies  under  A. 
panormis,  which  is  a  synonym  of  the  Burmese  species  A.  taras  (Doherty). 

The  species  is  purely  Sundanian,  not  extending  to  Palawan  or  the  islands  off  the  west  coast  of  Sumatra, 
and,  at  least  in  the  Malay  Peninsula,  flies  at  higher  average  elevations  than  the  other  species  of  the 
subgenus,  being  seldom  found  below  800  m. 

I  recognise  three  weak  subspecies  based  on  a  mean  of  differences. 

Key  to  the  subspecies  of  A.  (F.)  portunus 

1  cf  underside  of  forewing  with  blackish  submarginal  spots  normally  absent 2 

-     cf  underside  of  forewing  with  blackish  submarginal  spots  normally  present...     portunus  pyxus  (p.  30) 

2  cf  underside  of  forewing  with  postdiscal  series  ill-marked.  $  underside  usually  pale  greyish  buff, 

with  forewing  apex  shaded  with  ochreous portunus  portunus  (p.  29) 

Cf  underside  of  forewing  with  postdiscal  series  narrow  but  usually  clearly  defined.  $  underside 
greyish  white  with  forewing  apex  only  narrowly  and  faintly  shaded portunus  maitus  (p.  29) 

Allotinus  (Fabitaras)  portunus  portunus  (de  Niceville) 

Paragerydus  portunus  de  Niceville,  1894:  27,  pi.  5,  fig.  14  cf .  Syntypes,  JAVA  (?ZSI). 

Allotinus  taras  narsares  Fruhstorfer,  1913:  370;  1916:  813;  Corbet,  19396:  74,  partim.  LECTOTYPE  $, 

JAVA  (BMNH),  here  designated  [examined].  Syn.  n. 

Allotinus  portunus  portunus  (de  Niceville)  Fruhstorfer,  1914:  22;  1916:  813. 
[Allotinus  strigatus  dositheus  Fruhstorfer,  1914:  22  partim,  $  nee  cf .] 
[Allotinus  taras  (Doherty);  Piepers  &  Snellen,  1918:  14,  pi.  19,  fig.  16  $.  Misidentification.] 
Allotinus  portunus  (de  Niceville);  Piepers  &  Snellen,  1918:  15,  pi.  20,  figs  18a  cf ,  18b  $. 
Allotinus  panormis  portunus  (de  Niceville);  Corbet,  19396:  75. 

On  the  underside  the  ground  colour  is  variable  in  both  sexes.  In  the  male  it  varies  from  pale  buff  to  rufous 
buff,  and  generally  the  postdiscal  markings  are  ill-defined  and  may  be  absent  as  in  the  example  figured  by 
de  Niceville.  In  the  female  the  ground  colour  may  be  as  buff  as  in  the  male,  but  in  some  examples,  as  in  the 
lectotype  of  narsares,  which  apparently  was  the  model  for  the  figure  of  A.  taras  in  Piepers  &  Snellen  (1918), 
it  is  pale  greyish.  The  blackish  submarginal  spots  on  the  underside  of  the  forewing  are  usually  missing  in  the 
male  but  present  in  the  female  and  at  most  only  weakly  edged  with  white. 

I  designate  as  lectotype  of  narsares  a  female  in  BMNH  labelled  /Type  [red]/Java  Occident.  Sukabumi 
2000'  ex  coll.  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/narsares  Frhst.  [in  Fruhstorfer's  hand]/. 

DISTRIBUTION.  Java. 

Allotinus  (Fabitaras)  portunus  maitus  Fruhstorfer 
(Fig.  19,  Cf  genitalia) 

Allotinus  portunus  maitus  Fruhstorfer,  1914:  21;  1916:  813.  LECTOTYPE  cf,  SUMATRA  (BMNH),  here 
designated  [examined]. 


30  J.  N.  ELIOT 


Fig.  19    Allotinus  (Fabitaras)  portunus  maitus  Fruhstorfer;  Malay  Peninsula.  Male  genitalia. 

Allotinus panormis  fruhstorferi  Corbet,  19396: 74,  fig.  7,  cf  genitalia;  Fleming,  1975:  22,  pi.  58,  fig.  L47  $ ; 

Eliot,  1978:  240.  Holotype  cf ,  WEST  MALAYSIA  (BMNH)  [examined].  Syn.  n. 
Allotinus  panormis  maitus  Fruhstorfer;  Corbet,  19396:  74. 

In  the  male  the  postdiscal  markings  on  the  underside  are  usually  more  clearly  defined  than  in  subsp. 
portunus  and  the  submarginal  blackish  spots  are  missing;  in  the  female  the  ground  colour  is  more  greyish 
white. 

I  designate  as  lectotype  of  maitus  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/CMB 
111.  94/Fruhstorfer  Coll.  B.M.  1933-131/portunus  maitus  Frhst.  [in  Fruhstorfer's  hand]/.  A  female  labelled 
/Type  [red]/Type  [Fruhstorfer  orange]/Sumatra  Monies  Battak  ex  coll.  H.  Fruhstorfer/Fruhstorfer  Coll. 
B.M.  1933-131/allotype  Allotinus  portunus  maitus  Fruh.  [in  Corbet's  hand]/  is  a  paralectotype. 

DISTRIBUTION.  Sumatra;  West  Malaysia. 

Allotinus  (Fabitaras)  portunus  pyxus  (de  Niceville) 

Paragerydus pyxus  de  Niceville,  1894:  27,  pi.  5,  fig.  2  cf .  Holotype  cf ,  BORNEO:  Kina  Balu  (?  ZSI). 
Paragerydus  waterstradti  H.  H.  Druce,  1895:  562,  pi.  31,  figs  1  cf,  2  $.  LECTOTYPE  cf,  BORNEO 

(BMNH),  here  designated  [examined].  [Synonymised  by  Fruhstorfer,  1914:  22.] 
Paragerydus  waterstradti  ab.  absens  H.  H.  Druce,  1895:  562. 

Allotinus  portunus  pyxus  (de  Niceville);  Fruhstorfer,  1914:  22;  1916:  813,  pi.  141i,  cf  $. 
Allotinus  panormis  pyxus  (de  Niceville);  Corbet,  19396:  75. 

Females  appear  to  be  inseparable  from  the  nominate  subspecies  but  in  males  the  blackish  submarginal 
spots  on  the  underside  of  the  fore  wing  are  more  often  present. 

De  Niceville,  who  named  the  subspecies  from  a  single  male,  differentiated  it  from  subsp.  portunus  by  the 
more  rufous  tone  of  the  upperside  and  the  pale  rufous  instead  of  pale  ochreous  colour  of  the  underside. 
However,  the  ground  colour  is  very  variable  in  all  three  subspecies  and  is  not  a  reliable  character.  De 
Niceville's  figure  is  misleading,  as  it  does  not  show  the  visual  brand  which  is  mentioned  in  his  description. 

H.  H.  Druce  described  waterstradti  from  syntypes  in  coll.  Staudinger  and  in  his  own  collection.  I 
designate  as  lectotype  a  male  in  BMNH  labelled  /Kina  Balu  Waterstr./P.  waterstradti  co-type  H.  H. 
Druce/ex  coll.  Hamilton  Druce/Joicey  Bequest  Brit.  Mus.  1934-120/.  As  there  is  no  specimen  in  BMNH  of 
ab.  absens  H.  H.  Druce,  it  is  likely  that  the  name  applies  to  a  specimen  in  coll.  Staudinger. 

DISTRIBUTION.  Borneo,  apparently  only  known  from  Mt  Kina  Balu. 

Subgenus  PARAGERYDUS  Distant 

Paragerydus  Distant,  1884:  207.  Type-species:  Miletus  horsfieldi  Moore,  1857:  19,  pi.  la,  fig.  1,  by 

designation  of  Kirby,  [1885]:  191. 
Miletographa  Rober,  1892:  277.  Type-species:  Miletus  drumila  Moore,  [1866]:  777,  pi.  41,  fig.  12,  by 

monotypy.  [Synonymised  by  Fruhstorfer,  1913:  371.] 


THE  MILETINI  31 

Antenna  a  little  over  half  length  of  costa,  slightly  longer  than  in  subgen.  Allotinus  and  shorter  than  in 
subgen.  Fabitaras.  Antennal  segments  number  about  40  to  60.  Shaft  segments  of  the  smaller  species 
relatively  longer,  consequently  fewer  in  number;  thus  in  the  nivalis-group  and  in  A.  corbeti  there  are  about 
40  segments,  whereas  in  the  larger  species,  such  as  A.  horsfieldi  and  A.  apries,  there  are  about  58  segments. 
Species  of  intermediate  size  have  an  intermediate  number  of  segments,  for  example  in  A.  unlcolor  there 
are  usually  about  45.  In  dwarf  individuals,  occurring  most  frequently  in  A.  horsfieldi  and  A.  unicolor,  there 
is  no  diminution  in  the  number  of  segments.  Venation  shows  less  individual  variation  than  in  Fabitaras. 
Hindwing  without  humeral  vein.  Forewing  veins  MI  and  R5  usually  connate  or  briefly  stalked,  but  in  the 
nivalis-group  they  may  be  just  separate  at  their  origins.  In  males,  vein  M3  of  forewing  basally  swollen, 
clothed  with  small  specialised  scales,  those  of  the  nivalis-group  being  relatively  large.  In  males,  abdominal 
hair  tufts  prominent,  permanently  extruded  and,  in  the  genitalia,  the  valva  has  a  truncate  apex,  except  in 
A.  davidis,  and  its  ventral  edge  prolonged  into  a  more  or  less  pointed  process.  With  only  a  few  exceptions, 
the  interspecific  differences  in  the  valva  are  very  slight  and  perhaps  inconstant;  in  general  the  genitalia  give 
little  assistance  in  identification. 

Early  authors,  up  to  Fruhstorfer  (1913;  1916),  used  Paragerydus  in  a  wider  sense,  either  as  a  genus  or 
subgenus  (Artengruppe)  to  include  the  species  here  placed  in  Fabitaras. 

The  subgenus  ranges  from  north  India  to  Sundaland,  the  Lesser  Sunda  Is.,  Philippines  and  Sulawesi 
(including  Sula  Is.).  It  comprises  16  species. 

Key  to  the  species  of  subgenus  Paragerydus 

1  Underside  mottled  with  brown  specks  and  striae;  postdiscal  series  not  outlined  by  darker  lines 

nor  catenulate.  Smaller,  forewing 9-0-23-0  mm 2 

-  Underside  mottled  with  brown  spots  ringed  with  pale  buff;  postdiscal  series  catenulate  and 

outlined  by  darker  lines.  Larger,  forewing  usually  over  23-0  mm drum  Ha  (p.  56) 

2  Underside  of  forewing  without  a  white  fleck  at  end  of  vein  R5  (horsfieldi-group) 3 

-  Underside  of  forewing  with  a  white  fleck  at  end  of  vein  R5  (nivalis-group) 14 

3  Underside  with  postdiscal  series  more  or  less  the  same  size  throughout.  Upperside  of  hindwing 

unmarked  brown 4 

Underside  with  postdiscal  spots  below  vein  M3  on  forewing  much  larger  than  those  above,  and 
spots  above  vein  MI  on  hindwing  much  larger  than  those  below.  Upperside  of  hindwing 
partly  white  in  cf  $  or  in  $  alone 13 

4  Upperside  of  forewing  brown;  white  patch  or  streak  present  in  9  of  two  species.  Hindwing 

brown,  of  same  shade  as  forewing;  termen  crenulate  (except  in  A .  apries) 5 

Upperside  of  forewing  white  to  wing  base,  with  dark  brown  margin  and  costa.  Hindwing 
uniform  pale  buff-brown;  termen  not  crenulate parapus  (p.  51) 

5  $  upperside  brown,  cf  with  visual  brand,  if  present,  not  extending  into  cell  basad  of  origin  of 

vein  Cui 6 

-  $  upperside  of  forewing  with  white  patch  or  streak,  cf  (where  known)  with  visual  brand 

extending  obscurely  into  cell  basad  of  origin  of  vein  Cu2 samarensis  (p.  37) 

6  Underside  of  forewing  with  postdiscal  spots  in  spaces  4,  5  and  6  more  or  less  in  line  and 

equidistant ,  or  with  the  spot  in  space  5  nearer  to ,  and  often  touching,  the  spot  in  space  6 7 

Underside  of  forewing  with  postdiscal  spots  in  spaces  4,  5  and  6  on  an  uneven  curve,  that  in 
space  5  overlapping  that  in  space  4  and  well  separated  from  that  in  space  6  macassarensis  (p.  39) 

7  Forewing  veins  MI  and  R5  connate  or  with  a  common  stalk  not  more  than  1-0  mm  long. 

Underside  of  forewing  with  postdiscal  spot  in  space  2  nearer  termen  than  the  spots  in  spaces  3 
and  Ib  (if  present),  cf  $  with  hindwing  cilia  elongated  into  tufts  at  vein  endings,  termen 
crenulate  in  $ 

-  Forewing  veins  MI  and  R5  with  a  stalk  more  than  1-0  mm  long.  Underside  of  forewing  with 

postdiscal  spots  in  spaces  3,  2  and  Ib  (if  present)  in  line  and  parallel  to  termen.  cf  $  with 
hindwing  cilia  barely  longer  at  vein  endings  and  termen  not  crenulate  in  $  apries  (p.  42) 

8  cf  upperside  of  forewing  with  a  visual  brand ,  vein  M3  swollen  for  about  half  its  length  or  longer  9 
Cf  upperside  of  forewing  without  a  visual  brand,  vein  M3  swollen  for  only  one-quarter  of  its 

length. 
Small,  forewing  10-0-12-0 mm corbeti  (p.  44) 

9  Forewing  veins  Ml  and  R5  usually  with  a  very  short  common  stalk,  cf  valva  with  terminal 

process  not  rising  above  costa ;  tip  of  uncus  not ,  or  only  a  little ,  produced 10 

-  Forewing  veins  M1  and  R5  usually  connate,  cf  valva  with  terminal  process  curved  upwards, 

ending  just  above  costa;  tip  of  uncus  produced ,  rostriform horsfieldi  (p.  32) 


32  J.   N.  ELIOT 

10  cf  forewing  with  vein  A/3  swollen  for  at  least  three-fifths  of  its  length;  visual  brand  comparative- 

ly wide 11 

-  cf  forewing  with  vein  M3  swollen  for  only  half  its  length  or  slightly  over;  visual  brand 
comparatively  narrow,  less  than  1-5  mm  wide  (except  in  subspp.  continentalis ,  and  moorei 
sometimes,  of  A.  unlcolor) 12 

11  cf  valva  with  terminal  process  comparatively  long  and  narrow  (Fig.  21).  Underside  of 

sympatric  taxa  greyish  white;  on  hindwing  postdiscal  spot  in  space  6  usually  more  or  less 

below  that  in  space  7 leogoron  (p.  34) 

Cf  valva  with  terminal  process  short  and  broad  (Fig.  22).  Underside  ground  colour  very  pale 
buff;  on  hindwing  postdiscal  spot  in  space  6  nearly  always  inside  the  spot  in  space  7  and 
sometimes  nearly  mid-way  to  the  end-cell  bar melos  (p.  36) 

12  Underside  ground  colour  greyish  white  to  pale  buff.  Comparatively  small,  with  forewing 

10-5-18-0  mm.  cf  valva  (Figs  29,  30)  comparatively  broad,  with  terminal  process  in  centre 

line unicolor  (p.  45) 

Underside  chalky  white.  Larger,  forewing  18-0-20-0  mm.  cf  valva  narrower  (Fig.  31),  with 
terminal  process  nearly  in  line  with  ventral  edge paetus  (p.  50) 

13  cf  upperside  greyish  brown.  $  with  a  whitish  discal  patch  on  forewing  and  an  obscure  whitish 

streak  in  space  5  on  hindwing.  cf  9  discocellular  veins  on  hindwing  not  darkend 

luzonensis  (p.  40) 
Cf  $  upperside  partly  white  on  both  wings.  Discocellular  veins  blackened albatus  (p.  41) 

14  Underside  of  hindwing  with  central  spot  in  space  7  at  least  partly  blackened.  Smaller,  forewing 

9-0-14-5  mm 15 

Underside  of  hindwing  with  central  spot  in  space  7  not  blackened  (may  be  darker  brown  than 
other  spots).  Larger,  forewing  12-5-15-5  mm nivalis  (p.  51) 

15  cf  valva  with  costa  incised  shortly  before  apex  (as  in  all  preceding  species).  Underside  of 

hindwing  with  postdiscal  spot  in  space  7  not  blackened  on  its  inner  edge,  except  sometimes  in 
the  dry  season  form  in  Burma;  ground  colour  greyish  white,  except  in  Philippines  where  it  is 

pale  buff substrigosus  (p.  53) 

Cf  valva  with  costa  entire.  Underside  of  hindwing  with  postdiscal  spot  in  space  7  blackened  on 
its  inner  edge;  ground  colour  pale  buff.  Not  found  in  Burma  or  Philippines davidis  (p.  55) 

Allotinus  (Paragerydus)  hors field!  (Moore) 

(Fig.  20,  cf  genitalia) 
Miletus  horsfieldi  Moore,  1857:  19. 

This  and  three  succeeding  species,  leogoron,  melos  and  macassarensis ,  form  a  confusing  group  difficult  to 
separate  by  superficial  characters.  In  the  males  of  all  four  species  the  swelling  of  vein  M3  and  the  visual 
brand  on  the  forewing  vary  according  to  locality.  Venation  is  individually  variable,  but  in  A.  horsfieldi  veins 
MI  and  R5  of  the  forewing  are  nearly  always  connate ,  whereas  in  the  other  three  species  they  usually  share  a 
very  short  common  stalk.  In  both  sexes  the  hindwing  cilia  are  elongated  into  short  tufts  at  the  vein  endings 
and  females  have  the  hindwing  termen  crenulate.  In  the  female  of  A.  horsfieldi  the  hindwing  is  particularly 
strongly  crenulate.  In  A.  leogoron  the  crenulations  are  much  less  pronounced,  while  in  A.  melos  and  A. 
macassarensis  they  are  intermediate.  On  the  underside  the  markings  show  much  individual  variability  in 
their  density  and  in  the  position  of  the  postdiscal  series.  On  the  forewing  this  series  is  dislocated  at  vein  M3, 
with  the  stria  in  space  3  moved  basad  and  often  forming  with  the  stria  in  space  2  an  irregular,  oblique  stripe; 
but  the  degree  of  dislocation  is  variable  and  usually  slight  in  A.  macassarensis.  In  addition  the  stria  in  space 
Ib,  if  present,  is  moved  basad  in  relation  to  that  in  space  2. 

Males  of  A.  horsfieldi  can  always  be  identified  with  certainty  by  the  genitalia  (Fig.  20),  wherein  the 
ventral  tip  of  the  uncus  is  much  produced  and  the  terminal  process  of  the  valva  is  long  and  curved  up  above 
the  costa. 

In  both  sexes  A.  horsfieldi  shows  great  variability  in  size.  The  smaller  males,  with  forewing  length  as  little 
as  14  mm,  have  the  forewing  apex  and  termen  more  rounded  and  the  visual  brand  relatively  narrow,  but 
they  are  connected  by  a  complete  range  of  intermediates  to  the  largest  males,  with  forewing  length  up  to  23 
mm,  which  have  the  forewing  apex  more  pointed  and  the  termen  straighter. 

A.  horsfieldi  has  a  restricted  distribution  confined  to  Sundaland  excluding  Palawan.  Where  it  occurs  it  is 
usually  the  commonest  species  of  the  subgenus  apart  from  A.  unicolor. 


THE  MILETINI 


33 


Key  to  the  subspecies  of  A.  (P.)  horsfieldi 

1  Underside  pale  buff,  postdiscal  series  not  quadrate 2 

-  Underside  greyish  white,  postdiscal  series  comprising  large,  dark,  quadrate  spots 

horsfieldi  siporanus  (p.  34) 

2  Upperside  brown  without  a  reddish  tinge .  Underside  usually  lightly  marked 3 

-  Upperside  brown  with  a  reddish  tinge ,  especially  strong  in  9  and  in  basal  half  of  forewing  in  cf . 

Underside  more  strongly  marked horsfieldi  permagnus  (p.  33) 

3  cf  with  visual  brand  broad,  for  about  half  its  length  contiguous  with  vein  Cu\.  9  with  forewing 

disc  not  conspicuously  paler horsfieldi  horsfieldi  (p.  33) 

-  cf  with  visual  brand  narrower,  only  its  basal  quarter  touching  vein  Cu\.  9  with  forewing  disc 

conspicuously  paler horsfieldi  satelliticus  (p.  34) 

Allotinus  (Paragerydus)  horsfieldi  horsfieldi  (Moore) 

Miletus  horsfieldi  Moore,  1857:  19,  pi.  la,  fig.  2  cf .  LECTOTYPE  cf ,  JAVA  (BMNH),  here  designated 

[examined]. 

Allotinus  horsfieldi  horsfieldi  (Moore)  Fruhstorfer,  1913:  367;  1916:  812;  Corbet,  1939ft:  73. 
Allotinus  horsfieldii  [sic]  (Moore);  Piepers  &  Snellen,  1918: 12,  pi.  19,  figs  12a  Cf ,  12b  9, 14a  cf ,  14b  $. 

In  the  male  the  visual  brand  is  broad,  and  touches  vein  Cu^  for  about  half  its  length.  The  female  is  dull 
brown  without  a  reddish  tinge,  and  the  forewing  disc  is  only  a  little  paler.  On  the  underside,  especially  in 
females,  the  usual  lycaenid  markings  are  generally  light  and  may  be  faded  out,  as  in  the  extremes  shown  in 
Piepers  &  Snellen  (1918:  pi.  19,  figs  14a,  14b),  and  on  the  hindwing  the  postdiscal  spot  in  space  6  is  usually 
placed  well  inside  the  spot  in  space  7. 

I  designate  as  lectotype  a  male  in  BMNH  labelled  /Type  [red]/60-15  E. E.G. /Miletus  horsfieldi  cf 
M/GENITALIA  Slide  No  ASC  23  Allotinus/.  A  female  labelled  /Type  [red]/60-15  E.E.C./  is  a  paralecto- 
type. 

DISTRIBUTION.  Java. 

Allotinus  (Paragerydus)  horsfieldi  permagnus  Fruhstorfer 

(Fig.  20,  cf  genitalia) 
Paragerydus  horsfieldi  (Moore);  Distant,  1884:  207,  pi.  20,  fig.  7  '9'  recte  cf . 


Fig.  20    Allotinus  (Paragerydus)  horsfieldi  permagnus  Fruhstorfer;  Rhio  Archipelago:  Great  Karimon  I. 
Male  genitalia. 


34  J.  N.  ELIOT 

Allotinus  horsfieldi permagnus  Fruhstorfer,  1913:  366;  1916:  812.  LECTOTYPE  d",  SUMATRA  (BMNH), 

here  designated  [examined]. 
Allotinus  horsfieldi  nessus  Corbet,  19396:  72,  fig.  15,  cf  genitalia;  Eliot,  1978:  240.  Holotype  cf ,  WEST 

MALAYSIA  (BMNH)  [examined].  [Synonymised  by  Eliot,  1967:  66.] 
[Allotinus  leogoron  lindus  Corbet,  19396:  73,  partim  $  nee  d*.  Misidentification.] 
Allotinus  horsfieldi  permagnus  Fruhstorfer;  Eliot,  1967:  66,  fig.  1,  cf  genitalia;  Fleming,  1975:  21,  pi.  58, 

fig.  L43  O". 

The  subspecies  is  best  distinguished  by  the  female  which  is  reddish  brown  with  the  forewing  disc  only  very 
slightly  paler.  The  female  named  by  Fruhstorfer  as  f .  infumata  is  well  within  the  normal  range  of  individual 
variation.  In  the  male  the  visual  brand  is  comparatively  narrow,  only  touching  vein  Cui  at  its  base,  and  the 
wing  base  of  the  forewing  has  a  more  reddish  tinge  than  the  darker  apical  region. 

I  designate  as  lectotype  of  permagnus  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer 
orange]/W.  Sumatra  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/horsfieldi  permagnus  Fruhst.  [in 
Fruhstorfer's  hand]/.  The  specimen,  which  is  rather  more  lightly  marked  on  the  underside  than  usual,  has 
lost  its  abdomen.  A  female  labelled  /Type  [red]/Sumatra  Montes  Battak  ex  coll.  H.  Fruhstorfer/^ 
horsfieldi  Selesseh  19.  xii.  94/Fruhstorfer  Coll.  B.M.  1933-131/horsfieldi  permagnus  Fruhst.  [in  Fruhstor- 
fer's hand]/  is  a  paralectotype. 

I  designate  as  lectotype  of  infumata  a  female  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer 
orange]/N.  Oekor  viii.  94/Fruhstorfer  Coll.  B.M.  1933-131/infumata  Frhst.  [in  Fruhstorfer's  hand]/. 

DISTRIBUTION.  Malay  Peninsula,  including  Singapore;  Sumatra;  Bangka;  Borneo;  Peninsular  Thailand 
(Pinratana).  Three  females  in  BMNH  from  Batu  Is.  are  provisionally  included  in  this  subspecies;  they  are 
small  with  the  underside  ground  colour  whiter  and  the  postdiscal  markings  rather  dark  and  heavy,  showing 
an  approach  to  subsp.  siporanus.  They  are  also  rather  similar  to  A.  leogoron  batuensis,  with  which  Eliot 
(1967)  originally  confused  them  and  from  which  they  can  be  distinguished  by  their  more  crenulate 
hindwingtermen. 

Allotinus  (Paragerydus)  horsfieldi  siporanus  Riley 

Allotinus  horsfieldi siporana  Riley,  1944:  253.  Holotype  $.  MENTAWAI  Is.:  Sipora  (BMNH)  [examined]. 

The  subspecies  was  described  from  a  single  large  female  in  extremely  battered  condition;  the  whole  of  the 
outer  half  of  the  left  hindwing  and  the  tornal  quarter  of  the  right  hindwing  below  vein  Cu\  are  missing.  The 
greyish  white  ground  colour  of  the  underside  and  heavy  dark  brown  markings  suggest  that  it  might  pertain 
to  A.  leogoron,  but  such  crenulations  as  remain  on  the  termen  of  the  right  hindwing  suggest  A.  horsfieldi, 
and  it  seems  best  to  leave  Riley's  combination  unchanged  pending  the  discovery  of  the  male. 

DISTRIBUTION.  Mentawai  Is. 

Allotinus  (Paragerydus)  horsfieldi  satelliticus  Fruhstorfer 

Allotinus  horsfieldi  satelliticus  Fruhstorfer,  1913:  366;  1916:  812;  Eliot,  1967:  66.  LECTOTYPE  $, 
ENGANO  I.  (BMNH),  here  designated  [examined]. 

In  the  male  the  visual  brand  is  like  that  of  subsp.  permagnus,  but  in  other  respects  the  subspecies  more 
nearly  resembles  the  nominate  subspecies,  especially  in  the  female  which  is  without  a  reddish  tinge  on  the 
upperside.  However,  this  sex  differs  from  Javanese  females  by  having  a  very  prominent  paler  discal  patch 
on  the  forewing. 

I  designate  as  lectotype  a  female  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/Engano 
April-Juli  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/horsfieldi  satelliticus  Frhst.  [in  Fruhstorfer's 
hand]/. 

DISTRIBUTION.  Engano  I. 

Allotinus  (Paragerydus)  leogoron  Fruhstorfer 

(Fig.  21,  cf  genitalia) 
Allotinus  leogoron  Fruhstorfer,  1916:  811. 

This  species  has  been  much  confused  in  the  past  with  A.  horsfieldi,  from  which  the  male  is  readily  separated 
by  the  genitalia,  wherein  the  ventral  tip  of  the  uncus  is  not  produced  and  the  terminal  process  of  the  valva  is 


THE  MILETINI 


35 


fairly  long  and  narrow  but  not  upturned.  Females  are  best  distinguished  by  the  hindwing  termen,  which  is 
less  crenulate  than  in  horsfieldi,  but  the  cilia  bear  slightly  longer  and  narrower  tufts  at  the  vein  endings.  On 
the  forewing  veins  MI  and  R5  usually  have  a  short  common  stalk  and,  except  in  Java,  the  ground  colour  of 
the  underside  is  whiter  with  darker,  more  contrasted  markings  than  in  horsfieldi. 
A.  leogoron  has  the  same  distribution  as  A.  horsfieldi.  I  recognise  four  subspecies. 

Key  to  the  subspecies  of  A.  (P.)  leogoron 

1  cf  visual  brand  narrow,  only  its  base  touching  vein  C«i  2 

-  cf  visual  brand  wide,  touching  vein  Cu\  throughout  half  its  length leogoron  leogoron  (p.  35) 

2  Underside  greyish  white  with  dark  markings 3 

-  Underside  pale  buff  with  markings  smaller  and  paler  brown leogoron  plessis  (p.  36) 

3  d"  visual  brand  with  only  its  extreme  base  touching  vein  Cu\.  Underside  of  hindwing  with 

postdiscal  spot  in  space  6  below  that  in  space  7,  as  in  the  nominate  subspecies 

leogoron  nor  muni  (p.  36) 

-  cf  visual  brand  a  little  wider,  touching  vein  Cu\  for  a  quarter  of  its  length .  Underside  of  hindwing 

with  postdiscal  spot  in  space  6  placed  inside  that  in  space  7,  sometimes  almost  half-way  to 
cell-end  bar leogoron  batuensis  (p.  36) 

Allotinus  (Paragerydus)  leogoron  leogoron  Fruhstorfer 

(Fig.  21,  cf  genitalia) 

Allotinus  horsfieldi permagnus  $-f.  intricata  Fruhstorfer,  1913:  366;  1916:  812.  LECTOTYPE  $,  SUMATRA 

(BMNH),  here  designated  [examined].  [Unavailable  name.]  [Synonymised  by  Eliot,  1967:  68.] 
Allotinus  leogoron  Fruhstorfer,  1916:  811.  LECTOTYPE  cf,  SUMATRA  (BMNH),  here  designated 

[examined]. 
Allotinus  continental  vadosus  Corbet,  19396:  72.  Holotype  cf ,  WEST  MALAYSIA  (BMNH)  [examined]. 

[Synonymised  by  Eliot,  1967:  69.] 
Allotinus  leogoron  leogoron  Fruhstorfer;  Corbet,  19396:  73;  Eliot,  1967:  68,  fig.  5,  cf  genitalia;  1978:  240; 

Fleming,  1975:  21,  pi.  58,  fig.  L44  cf . 
Allotinus  leogoron  lindus  Corbet,  19396:  73,  partim  cf  nee  $.  Holotype  cf,  WEST  MALAYSIA  (BMNH) 

[examined].  [Synonymised  by  Eliot,  1967:  69.] 

The  male  can  be  separated  from  that  sex  of  sympatric  A.  horsfieldi  by  the  wider,  more  sharply  defined 
visual  brand  which  touches  vein  Cu\  throughout  its  basal  half,  and  the  female  by  the  less  crenulate  hindwing 
termen.  In  addition,  the  underside  is  whiter  and  the  markings  are  darker  brown. 


Fig.  21     Allotinus  (Paragerydus)  leogoron  leogoron  Fruhstorfer;  Malay  Peninsula.  Male  genitalia. 


36  J.  N.  ELIOT 

I  designate  as  lectotype  of  leogoron  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
CMB  viii.  94/A.  leogoron  Fr.  [in  Fruhstorfer's  hand]/  and  of  intricata  a  female  in  BMNH  labelled  /Type 
[red]/Type  [Fruhstorfer  orange]/Sumatra  Montes  Battak  ex  coll.  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M. 
1933-131/$  horsfieldi  CMB  viii.  94/intricata  [in  Fruhstorfer's  hand]/. 

DISTRIBUTION.  Malay  Peninsula;  Sumatra;  Bangka;  Peninsular  Thailand  (Pinratana). 

Allotinus  (Paragerydus)  leogoron  normani  Eliot 
Allotinus  leogoron  normani  Eliot,  1967:  69.  Holotype  cf ,  BORNEO  (BMNH)  [examined]. 

Differs  from  the  nominate  subspecies  only  in  that  the  male  has  the  visual  brand  much  narrower,  only 
touching  vein  Cui  at  its  extreme  base. 

DISTRIBUTION.  Borneo. 

Allotinus  (Paragerydus)  leogoron  batuensis  Eliot 
Allotinus  leogoron  batuensis  Eliot,  1967:  69.  Holotype  cf ,  BATU  Is.  (BMNH)  [examined]. 

A  small  subspecies,  with  a  slightly  more  reddish  brown  tinge  than  in  the  two  preceding  subspecies.  In  the 
male  the  visual  brand  is  intermediate  between  that  of  leogoron  and  that  of  normani,  touching  vein  Cui 
along  its  basal  quarter.  It  differs  on  the  underside  of  the  hindwing  in  that  the  postdiscal  spot  in  space  6  is 
placed  inside  the  spot  in  space  7. 

DISTRIBUTION.  Batu  Is. 

Allotinus  (Paragerydus)  leogoron  plessis  Eliot 

Allotinus  leogoron  plessis  Eliot,  1967:  69.  Holotype  cf ,  JAVA  (BMNH)  [examined]. 

In  the  male  the  visual  brand  is  narrow,  as  in  subsp.  normani,  whereas  the  brand  of  sympatric  A  horsfieldi 
horsfieldi  is  as  wide  as  in  subsp.  leogoron.  The  subspecies  is  further  distinguished  by  the  pale  buff  underside 
on  which  the  markings  are  smaller  and  less  contrasted. 

DISTRIBUTION.  Java. 

Allotinus  (Paragerydus)  melos  (H.  H.  Druce) 

(Figs  22,  cf  genitalia;  64  cf ) 

[Paragerydus  horsfieldi  (Moore)  sensu  Semper,  1889:  164  partim,  pi.  31,  figs  19  cf,  20  $.  Misidentifica- 

tion.] 
Paragerydus  melos  H.  H.  Druce,  1896:  652.  LECTOTYPE  cf ,  PHILIPPINES:  Cagayan  Sulu  (BMNH),  here 

designated  [examined] 

Allotinus  melos  (Druce)  Fruhstorfer,  1913:  369. 
Allotinus  horsfieldi  leos  [sic]  (Druce);  Fruhstorfer,  1916:  812. 
Allotinus  horsfieldi  reverdini  Fruhstorfer,  1916:  812.  LECTOTYPE  cf ,  PHILIPPINES  (SM),  here  designated 

[examined].  Syn.  n. 


melos  (H.  H.  Druce);  Eliot,  1967:  68. 
re verdini  Fruhstorfer;  Eliot,  1967:  68. 
talu  Eliot,  1967:  68.  Holotype  cf ,  BORNEO:  Pulo  Laut  (BMNH)  [examined]. 


Allotinus  macassariensis  [sic 
Allotinus  macassariensis  [sic 
Allotinus  macassariensis  [sic 
Syn.  n. 

This  species  replaces  A.  horsfieldi,  with  which  it  has  been  confused  in  the  past,  in  the  southern  Philippines. 
Both  species  are  found  in  Borneo,  but  A.  melos  seems  to  be  the  commoner  throughout  most  of  the  island. 
Sympatric  examples  of  the  two  species  are  very  difficult  to  separate  by  external  characters,  but  males  may 
be  separated  by  the  genitalia  without  difficulty;  in  melos  the  terminal  process  of  the  valva  is  short  and  broad 
and  the  tip  of  the  uncus  is  not  produced.  Females  may  be  impossible  to  separate  with  certainty,  but  on  the 
underside  of  the  hindwing  of  melos  the  postdiscal  spot  in  space  6  is  usually  placed  inside  the  spot  in  space  7, 
sometimes  almost  half-way  to  the  end-cell  bar,  whereas  in  horsfieldi  these  spots  usually  overlap.  In  addition 
the  hindwing  termen  of  melos  is  very  slightly  less  crenulate. 

Eliot  (1967)  treated  melos  as  conspecific  with  the  allopatric  A.  macassarensis ,  but  in  view  of  constant 
small  differences  in  the  male  genitalia  and  in  the  arrangement  of  the  postdiscal  spots  in  spaces  4, 5  and  6  on 
the  underside  of  the  forewing  (see  p.  39),  I  no  longer  maintain  this  combination. 


THE  MILETINI 


37 


Fig.  22    Allotinus  (Paragerydus)  melos  (H.  H.  Druce);  Mindanao.  Male  genitalia. 


Males  have  vein  M3  swollen  for  three-fifths  of  its  length  and  the  visual  brand  is  clearly  defined  and  about 
1-75  mm  wide.  The  underside  shows  very  great  individual  variation  in  the  intensity  of  the  markings  and  in 
the  placing  of  the  postdiscal  series.  The  postdiscal  series  is  quite  well  separated  from  the  wing  margins  in 
typical  examples  but,  especially  in  north-east  Borneo,  examples  occur  in  which  this  series  is  placed  close  to 
the  termen,  as  in  A.  macassarensis .  On  the  hindwing  the  postdiscal  spot  in  space  6  is  usually  placed  inside 
that  in  space  7,  sometimes  as  much  as  mid-way  to  the  end-cell  bar,  and  this  character  may  be  of  assistance  in 
separating  Bornean  females  from  sympatric  A.  horsfieldi  permagnus . 

H.  H.  Druce  stated  that  the  types  of  melos  were  in  Mus.  Cator  and  Druce.  There  is  in  coll.  Cator  (now  in 
BMNH)  a  long  series  of  both  sexes,  none  of  which  has  been  labelled  type,  and  in  the  main  BMNH  coll.  a 
pair  ex  H.  H.  Druce  coll.  labelled  as  paratypes.  Of  the  latter  I  designate  the  male  as  lectotype;  it  is  labelled 
/Paratype  [yellow]/Cagayan  3.  6.  94/Paragerydus  melos  d"  co-type  H.  H.  Druce/ex  coll.  Hamilton  Druce 
1919/Joicey  Bequest  Brit.  Mus.  1934-120/.  The  female,  labelled  /Paratype  [yellow]/Cagayan  2.  6.  94/P. 
melos  $  co-type  H.  H.  Druce/ex  Coll.  Hamilton  Druce  1919/Joicey  Bequest  Brit.  Mus.  1934-120/,  as  well 
as  the  series  in  coll.  Cator,  are  paralectotypes. 

When  naming  reverdini,  Fruhstorfer  referred  to  large  males  in  coll.  Semper.  I  designate  as  lectotype  a 
male  in  SM  labelled /19/Bohol/298b/213/Coll.  C.  Semper/Original  of  Semper  PI.  31,  fig.  19  from  Mindanao. 
?  loc.  label  changed.  Det.  as  A.  melos  reverdini  Fruh.  C?  J.  N.  Eliot,  ix.  1982/.  The  male  figured  by  Semper 
on  pi.  31,  fig.  19  was  said  by  him  to  have  come  from  Mindanao.  I  think  it  more  likely  that  the  label  'Bohol' 
was  inadvertently  transferred  from  the  male  shown  in  fig.  18  (which  is  the  holotype  of  A.  posidion  georgius 
and  which  is  without  a  locality  label  -  see  p.  49)  than  that  Semper  made  a  mistake  in  the  legend  to  his  plate 
31. 

DISTRIBUTION.  Cagayan  Sulu;  Mindanao;  Palawan;  Balabac  I.;  Borneo  (many  localities,  including  Pulo 
Laut).  There  is  a  male  in  BMNH  labelled  /Sula  Mangoli,  Oct.  '97,  W.  Doherty/,  which  I  suspect  may  be 
wrongly  labelled  although  Doherty  certainly  collected  there  at  that  date.  Examples  from  Mindanao  tend  to 
be  rather  heavily  marked  on  the  underside.  Males  from  Palawan  and  Balabac  tend  to  have  the  visual  brand 
slightly  longer  and  narrower.  Examples  from  Pulo  Laut  are  slightly  richer  buff  on  the  underside  than 
typical  examples  from  Cagayan  Sulu,  but  the  difference  is  hardly  sufficient  to  justify  maintaining  talu  as  a 
distinct  subspecies.  There  is  in  coll.  Cator  a  series  of  seven  males  and  one  female  from  Melikop  and 
Sapagaya  (small  islands  off  the  north  coast  of  Sabah)  which  Cator  had  placed  separately  from  his  series  of 
melos  as  an  unrecognised  species;  they  are  small,  darker  brown  above  and  with  a  greyish  white  underside 
bearing  darker  markings.  Superficially  they  resemble  A.  leogoron  normani,  but  the  male  genitalia  prove 
them  to  be  A.  melos,  of  which  they  could  well  be  regarded  as  a  microsubspecies. 

Allotinus  (Paragerydus)  samarensis  sp.  n. 

(Figs  23,  cf  genitalia;  65,  66  $, 105  cf ) 

Upperside  of  both  sexes  blackish  brown,  almost  as  dark  as  the  shade  of  A.  macassarensis.  Male,  only 
known  from  the  nominate  subspecies,  distinctive  in  that  visual  brand  extends  obscurely  as  a  paler  streak 


38 


J.  N.  ELIOT 


Fig.  23    Allotinus  (Paragerydus)  samarensis  samarensis  sp.  n. ;  Samar.  Male  genitalia.  Phallus  with  juxta 
attached. 


into  fore  wing  cell  basad  of  origin  of  vein  Cu2.  Female  distinguished  by  the  possession  of  a  white  area  on 
forewing.  Underside  marked  like  A  melos. 

Male  genitalia  hardly  differ  from  those  of  A.  leogoron,  of  which  it  may  conceivably  be  a  subspecies; 
because  of  the  unusual  visual  brand  and  shorter  swelling  of  vein  M3  in  the  male,  and  the  distinctive 
white-marked  female,  I  think  it  is  best  regarded  as  a  distinct  species. 

The  species  is  only  known  from  a  very  few  examples  from  the  southern  Philippines  and  Sulawesi  in 
slightly  different  subspecies. 

Key  to  the  subspecies  of  A.  (P.)  samarensis 

1     Underside  pale  greyish  white.  $  upperside  of  forewing  with  white  area  extending  into  cell  well 

basad  of  origin  of  vein  Cu2 samarensis  samarensis  (p.  38) 

Underside  very  pale  buff.  $  upperside  of  forewing  with  white  area  not,  or  only  just,  entering  cell 

samarensis  russelli  (p.  38) 

Allotinus  (Paragerydus)  samarensis  samarensis  subsp.  n. 

(Figs  23,  cf  genitalia;  65  $,  105  cT) 

Cf  upperside  blackish  brown,  with  vein  M3  swollen  for  just  over  half  its  length,  visual  brand  about  1  -25  mm 
wide;  a  pale  streak  above  cubitus  extending  a  little  basad  of  origin  of  vein  Cu2  appears  to  be  an  extension  of 
the  brand.  Underside  marked  as  in  sympatric  A.  melos,  but  with  ground  colour  more  greyish  white. 

$  upperside  blackish  brown.  Forewing  with  a  creamy  white  streak  in  spaces  4  and  3  ending  2-5  mm  from 
termen  and  extending  into  cell  along  cubitus  to  beyond  origin  of  vein  Cu2.  Underside  similar  to  male. 

MATERIAL  EXAMINED 

Holotype  cf ,  Philippines:  Samar,  Bagacay,  2000',  31. v.  1979  (C.  G.  Treadaway)  (coll.  Treadaway). 

Paratypes.  Philippines:  1  $  (allotype),  west  Samar,  Hinabangan,  1000  m,  5.ii.l984  (coll.  Treadaway);  2 
Cf ,  data  as  holotype  (coll.  Treadaway). 

Excluded  from  type-series.  Mindanao:  1  $,  Surigao  Sur,  Tandag,  viii.  1981  (ex  Takanami  coll.) 
(BMNH),  with  the  white  streak  wider  and  extending  into  space  2. 

Allotinus  (Paragerydus)  samarensis  russelli  subsp.  n. 

(Fig.  66  $) 

Cf  unknown. 

$  forewing  19-0-21-0  mm.  Upperside  dark  brown  with  a  white  discal  patch  divided  by  dark  veins  on 
forewing,  measuring  4-0  mm  in  space  2  and  8-0  mm  in  space  3,  entering  base  of  space  4  and  lower  angle  of 
cell  where  it  is  sullied.  In  a  second  specimen  the  white  patch  is  smaller,  not  extending  above  vein  M3  nor 
entering  cell.  Underside  very  pale  buff  with  brown  markings  arranged  as  in  nominate  subspecies. 


THE  MILETINI 


39 


MATERIAL  EXAMINED 
Holotype  $?>  Sulawesi:  east-central,  north-west  of  Morowali,  Kabalo,  450  m,  10.iii.1980  (A.  Bedford 


Paratype.  1  $  ,  data  as  holotype  (coll.  Bedford  Russell). 

Allotinus  (Paragerydus)  macassarensis  (Holland) 

(Figs  24  cf  ,  genitalia;  67  cf  ,  68  £) 
Paragerydus  macassarensis  Holland,  1891:  70. 

The  species  name  has  been  consistently  misspelled  macassariensis  in  subsequent  literature. 

Previously  (Eliot,  1967)  I  united  this  species  and  A.  melos,  but  in  view  of  small  but  apparently  constant 
differences  in  facies  and  male  genitalia  it  seems  best  to  treat  them  as  separate  allopatric  species. 

The  upperside  in  both  sexes  is  slightly  blacker  brown  than  in  the  other  species  of  the  subgenus.  On  the 
underside  the  postdiscal  markings,  which  are  usually  heavy  and  dark  chocolate  brown  on  an  off-white 
ground,  are  placed  closer  to  the  termen  than  in  the  preceding  species.  A  distinguishing  character  is  that  on 
the  forewing  the  postdiscal  spots  in  spaces  4,  5  and  6  are  on  an  irregular  curve,  with  the  spot  in  space  5 
overlapping  that  in  space  4  and  well  separated  from  that  in  space  6.  In  the  male  genitalia  the  valvae  differ 
from  those  of  A.  melos  in  two  respects;  the  terminal  process  is  narrower  and  slightly  longer  from  whichever 
angle  it  is  viewed  and,  in  lateral  view,  there  is  a  distinct  concavity  (indicated  in  Fig.  24  by  an  arrow)  about 
two-thirds  from  the  base. 

The  species  is  confined  to  Sulawesi  and  its  satellite  islands  and  is  represented  by  two  subspecies. 

Key  to  the  subspecies  of  A.  (P.)  macassarensis 

1     cf  vein  M3  of  forewing  swollen  for  over  two-thirds  of  its  length,  visual  brand  usually  less  than  1 

mm  wide  ..................................................................      macassarensis  macassarensis  (p.  39) 

-    Cf  vein  M3  of  forewing  swollen  for  only  half  its  length,  visual  brand  more  than  1  mm  wide 

macassarensis  menadensis  (p.  40) 

Allotinus  (Paragerydus)  macassarensis  macassarensis  (Holland) 

(Figs  24,  cf  genitalia;  67  cf  ,  68  £) 

Paragerydus  macassarensis  Holland,  1891:  70,  pi.  4,  fig.  5  $  .  Holotype  $  ,  SULAWESI  (not  located,  probably 

in  CM). 
Allotinus  horsfieldi  macassariensis  [sic]  (Holland)  Fruhstorfer,  1913:  368;  1916:  812,  pi.  141h. 


Fig.  24    Allotinus  (Paragerydus)  macassarensis  macassarensis  (Holland);  Sulawesi.  Male  genitalia.  Lower 
left,  right  valva  enlarged. 


40  J.  N.  ELIOT 

Allotinus  unicolor  damodar  Fruhstorfer,  1913:  369;  1916:  811.  LECTOTYPE  cf ,  SULAWESI  (BMNH), 

here  designated  [examined].  [Synonymised  by  Eliot.  1967:  68.] 
Allotinus  macassariensis  [sic]  (Holland);  Corbet,  19396:  72,  fig.  14  cf  genitalia. 
Allotinus  macassariensis  macassariensis  [sic]  (Holland);  Eliot,  1967:  68. 

In  the  male  the  swelling  of  vein  A/3,  extending  just  over  two-thirds  of  its  length,  is  the  longest  in  the 
subgenus  and  the  visual  brand  is  typically  very  narrow.  But  in  east-central  Sulawesi  there  is  a  tendency  for 
the  brand  to  become  wider  and  less  well-defined;  in  one  extreme  example  in  coll.  Bedford  Russell  (Fig.  64) 
there  is  a  diffuse  lighter  patch  which  reaches  across  vein  Cu\  more  than  half-way  to  vein  Cu2.  The  female 
has  the  forewing  disc  at  most  only  slightly  paler. 

I  designate  as  lectotype  of  damodar  a  small  but  otherwise  normal  male  in  BMNH  labelled  /Type 
[red]/Type  [Fruhstorfer  orange]/S.  Celebes  Tonus  27.xi.06/Fruhstorfer  Coll.  B.M.  1933-131/unicolor 
damodar  Frhst.  [in  Fruhstorfer's  hand]/.  A  female  labelled  /Type  [red]/S.  Celebes  Samangi  17.xi.06/ 
Fruhstorfer  Coll.  B.M.  1933-13 1/damodar  Frhst.  [in  Fruhstorfer's  hand]/  is  a  paralectotype. 

DISTRIBUTION.  South  to  central  Sulawesi;  Banggai  I. 

Allotinus  (Paragerydus)  macassarensis  menadensis  Eliot 

Allotinus  macassariensis  [sic]  menadensis  Eliot,  1967:  68.  Holotype  cf,  SULAWESI:  north  (BMNH), 
[examined]. 

Differs  from  the  nominate  subspecies  in  that  the  swollen  portion  of  vein  M3  in  the  male  is  only  half  its  length 
and  the  visual  brand  is  quite  well-defined  and  from  1-25  to  2-0  mm  wide.  In  both  sexes  the  forewing  disc 
may  be  lightened  by  a  scattering  of  white  scales,  extreme  examples  showing  an  approach  to  A.  samarensis 
russelli. 

DISTRIBUTION.  North  Sulawesi  as  far  south  as  Paloe  Bay  (Lat.  0°  35'  S).  A  single  male  from  Bangka  I.  (off 
the  tip  of  the  Minahassa  Peninsula)  is  more  lightly  marked  beneath  and  the  ground  colour  has  a  buff  tint. 

Allotinus  (Paragerydus)  luzonensis  Eliot  stat.  n. 

(Figs  25,  cf  genitalia;  69  cf ,  70  9) 

Allotinus  macassariensis  [sic]  luzonensis  Eliot,  1967:  68,  fig.  3  cf  genitalia.  Holotype  cf,  PHILIPPINES: 
Luzon  (BMNH)  [examined]. 

The  male  is  greyish  brown  above;  on  the  forewing  vein  M3  is  swollen  for  half  its  length  and  the  visual  brand 
is  rather  diffuse.  The  underside  is  buff,  closely  striated,  and  the  postdiscal  markings  in  spaces  2  and  3  on  the 


Fig.  25    Allotinus  (Paragerydus)  luzonensis  Eliot;  Luzon.  Male  genitalia. 


THE  MILETINI  41 

forewing  and  6  and  7  on  the  hindwing  are  broad  and  rather  blotchy.  The  female  has  a  small  white  patch  on 
the  forewing  disc  and  an  obscure  whitish  streak  in  space  5  of  the  hindwing.  In  these  white  patches  it  shows 
an  approach  to  the  female  of  A.  albatus  mendax,  but  they  are  less  developed  than  in  that  species,  and  the 
discocellular  veins  on  the  hindwing  are  not  blackened. 

The  species  must  be  closely  related  to  A .  albatus  because  of  the  same  arrangement  of  the  markings  on  the 
underside.  The  male  genitalia  of  the  only  example  dissected  are  intermediate  between  those  of  A.  albatus 
and  A.  melos. 

DISTRIBUTION.  Luzon. 

Allotinus  (Paragerydus)  albatus  C.  &  R.  Felder 

(Figs  26,  cT  genitalia;  71  £,  72  c?,  73  £) 
Allotinus  albatus  C.  &  R.  Felder,  [1865]:  287. 

The  species  is  readily  recognised  by  the  presence  in  both  sexes  of  white  areas  on  both  wings  and  by  the 
heavily  blackened  discocellular  veins  on  the  hindwing.  The  underside  is  white  marked  more  or  less  as  in  A. 
luzonensis,  and  like  this  species  veins  MI  and  R5  of  the  forewing  usually  have  a  very  short  common  stalk. 
The  male  genitalia  most  nearly  resemble  those  of  A  leogoron,  with  the  terminal  process  of  the  valva  rather 
longhand  slender. 
The  species  appears  to  be  very  rare  and  to  be  confined  to  Wallacea. 

Key  to  the  subspecies  of  A.  (P.)  albatus 

1  $  upperside  with  larger  white  areas,  on  forewing  extending  from  space  Ib  to  base  of  space  5,  on 

hindwing  from  outer  part  of  space  6  to  dorsum albatus  albatus  (p.  41) 

-  C?  $  upperside  with  smaller  white  areas,  on  forewing  not  below  mid-space  Ib  and  not  entering 

spaces  4  and  5,  on  hindwing  usually  not  below  vein  4 albatus  mendax  (p.  41) 

Allotinus  (Paragerydus)  albatus  albatus  C.  &  R.  Felder 

(Fig.  71$) 

Allotinus  albatus  C.  &  R.  Felder,  [1865]:  267.  Holotype  $,  SULAWESI  (BMNH)  [examined]. 
[Allotinus  fallax  major  Felder  f.  albatus  Felder  (=  maximus  Staudinger)  sensu  Fruhstorfer,  1913:  343, 
partim;  1916:  809,  partim,  asalbadus  [sic].] 

The  subspecies  is  known  only  from  the  female  holotype,  which  the  authors  stated  was  from  'Celebes 
Lorquin'.  It  was  therefore  surprising  to  find  the  holotype  bearing  labels  reading  /Halmaheira  Lorquin 
[round  blue]/Allotinus  albatus  Feld./Type  [red]/FELDER  COLLn. /albatus  n./Rothschild  Bequest  B.M. 
1939-1/.  As  no  Allotinus  species  is  known  from  east  of  Weber's  Line  I  feel  confident  that  the  locality 
'Halmaheira'  is  the  result  of  a  wrongly  acquired  label  and  that  the  specimen  came  from  north  Sulawesi, 
where  Lorquin  is  known  to  have  collected. 

The  forewing  is  white  dusted  with  brown  basally  and  along  the  dorsum  below  vein  A\,  and  with  an 
irregular  dark  brown  border  narrowest  at  vein  Cu2,  where  it  measures  2-5  mm;  above  vein  Cu-i  it  curves  to 
the  wing  base  through  the  upper  third  of  the  cell.  The  hindwing  is  white,  becoming  sullied  towards  the 
dorsum,  except  for  most  of  the  cell  and  spaces  8,  7  and  the  basal  half  of  space  6  which  are  brown;  the 
discocellular  veins  are  heavily  blackened.  The  underside  is  white  with  heavy  brown  markings. 

Fruhstorfer  confused  A.  albatus  with  large  examples  of  A.  major  with  extensive  white  areas  on  the 
forewing  which  were  named  f .  maximus  by  Staudinger. 

DISTRIBUTION.  Sulawesi. 

Allotinus  (Paragerydus)  albatus  mendax  subsp.  n. 

(Figs  26,  c?  genitalia;  72  a",  73  $) 

[Allotinus  fallax  C.  &  R.  Felder,  1865:  285,  partim,  pi.  35,  figs  25,  26  $;  Semper,  1889:  163,  partim. 
Misidentifi  cations .  ] 

Cf  forewing  length  20-0  mm.  Upperside  blackish  brown.  Forewing  with  a  discal  white  patch  7-0  mm  wide  at 
the  bases  of  spaces  3,  2  and  upper  part  of  Ib  and  entering  the  lower  angle  of  the  cell;  the  inner  and  lower 
part  of  the  cell  and  lower  part  of  space  Ib  paler  brown  than  the  marginal  and  costal  border.  Vein  M3  swollen 


42 


J.   N.   ELIOT 


:  .OP 


Fig.  26    Allotinus  (Paragerydus)  albatus  mendax  subsp.  n.;  Luzon.  Male  genitalia.  Lower  left,  internal 
view  of  left  valva  enlarged;  lower  centre,  lateral  view  of  phallus. 

for  just  under  three-fifths  of  its  length.  Hindwing  with  a  white  streak  filling  most  of  space  5,  the  upper  basal 
part  of  space  4  and  just  entering  the  cell;  discocellular  veins  heavily  blackened.  Underside  white,  with 
reddish  brown  markings  arranged  as  in  sympatric  A  luzonensis. 

9  similar  to  the  male,  except  that  the  white  streak  on  the  hindwing  is  wider  and  enters  space  6. 

MATERIAL  EXAMINED 

Holotype  cf,  Philippines:  Luzon,  bearing  labels  /Allotinus  mendax  Bd.  Manille/Ex  Musaeo  Dris 
BOISDUVAL/ex  Oberthur  Coll.  Brit.  Mus.  1927-3/ Allotinus  albatus  Feld.  [in  Corbet's  hand]/. 

Paratypes.  Philippines:  1  $  (allotype)  Luzon,  labelled  /N.  Luzon,  Whitehead.  94/Rothschild  Bequest 
B.M.  1939-1/  (BMNH);  1  cf ,  Luzon,  Bicol  Nat.  Park,  29.viii.1980  (Y.  Takanami)  (BMNH);  1  cf ,  Luzon, 
Quezon  Nat.  Park,  27.iv.1983  (Y.  Takanami)  (BMNH);  1  cf,  Luzon,  Banahao  Ridge,  v.1982  (A. 
Ballantine)  (coll.  Ballantine);  1  <j>,  Luzon,  Quezon  Nat.  Park,  1000',  Altimonan  Rd,  20.vi.1954  (coll. 
Treadaway);  1  $,  Quezon  Nat.  Park,  Altimonan  area,  28. iv.  1969  (coll.  Treadaway). 

Excluded  from  type-series.  Marinduque:  1  $ ,  Nu  Boac,  xi.1980  (coll.  Treadaway)  with  the  white  area  on 
the  hindwing  more  extensive  and  nearly  reaching  the  dorsum  and  the  discocellular  veins  less  heavily 
blackened.  Samar:  1  $,  east,  Borongan,  100',  10.viii.1979  (coll.  Treadaway),  resembling  the  allotype 
above,  but  beneath  with  the  postdiscal  markings  diffuse  and  obscure. 

Allotinus  (Paragerydus)  apries  Fruhstorfer 

(Figs  27  Cf  genitalia;  74, 106  cf ) 
Allotinus  horsfieldi  apries  Fruhstorfer,  1913:  344,  partim. 

Fruhstorfer 's  type-series  probably  comprised  more  than  one  species;  in  addition  he  confused  apries  with  A. 
strigatus.  I  use  apries  here  for  the  species  identified  as  such  by  Corbet  (19396). 

The  species  has  several  characters  which  enable  it  to  be  recognised  with  comparative  ease.  On  the 
forewing  veins  Ma  and  R5  share  a  long  common  stalk  averaging  1-5  mm.  On  the  underside  of  the  forewing 
the  postdiscal  series  is  not,  or  only  very  slightly,  dislocated  at  vein  M3  and  the  spots  in  spaces  4,3,2  and  Ib 
(if  present)  are  small,  rounded  and  well  separated  from,  and  parallel  to,  the  termen.  In  the  female  the 
hindwing  termen  is  almost  evenly  rounded  and  the  cilia  are  a  little  longer,  but  not  tufted,  at  the  vein 
endings.  In  the  male  genitalia  the  comparatively  slender  phallus  is  distinctive. 

The  species  occurs  throughout  Sundaland. 

Key  to  the  subspecies  of  A.  (P.)  apries 

1  Underside  pale  buff,  tending  to  become  darker  towards  forewing  apex  in  cf,  with  darker 

buff-brown  markings,  cf  visual  brand  2-0  mm  wide,  with  half  its  lower  edge  touching  vein  Cui  2 

Underside  greyish  white,  with  darker  greyish  brown  markings,  cf  visual  brand  1-5  mm  wide, 
with  only  basal  fifth  of  its  lower  edge  touching  vein  Cui  apries  ristus  (p.  44) 

2  Upperside  reddish  brown apries  apries  (p.  43) 

-    Upperside  brown  without  a  reddish  tint apries  dositheus  (p.  44) 


THE  MILETINI 


43 


Allotinus  (Paragerydus)  apries  apries  Fruhstorfer 

(Figs  27,  Cf  genitalia;  74  cf ) 

[Allotinus  horsfieldi  (Moore)  sensu  Swinhoe,  1910:  198,  partim,  pi.  617,  fig.  1  cf .  Misidentification.] 
Allotinus  horsfieldi  apries  Fruhstorfer,  1913:  344,  partim;  1916:  812,  partim,  pi.  141g  cf  nee  $.  LECTO- 

TYPE  cf ,  BORNEO  (BMNH),  here  designated  [examined]. 
Allotinus  strigatus  eupalion  Fruhstorfer,  1914:  22;  1916:  813.  LECTOTYPE  cf ,  SUMATRA  (BMNH),  here 

designated  [examined].  Syn.  n. 
Allotinus  apries  apries  Fruhstorfer;  Corbet,  1939ft:  70. 
Allotinus  apries  eupalion  Fruhstorfer;  Corbet,  1939ft:  70;  Fleming,  1975: 21,  pi.  58,  fig.  L42  cf ;  Eliot,  1978: 

240. 

In  both  sexes  the  underside  is  nearly  always  pale  buff,  with  darker  buff-brown  markings,  but  occasionally  it 
is  more  greyish  white  with  only  a  slight  buff  tint.  In  the  female  the  postdiscal  series  is  lightly  marked  and 
may  be  obsolete  on  the  forewing.  The  male  has  vein  A/3  swollen  for  a  little  over  three-fifths  its  length,  and 
the  visual  brand  is  clearly  defined  and  2-0  mm  wide.  The  female  is  more  reddish  brown  above,  with  the 
forewing  disc  a  little  paler. 

Fruhstorfer  described  A.  horsfieldi  apries  from  several  males  from  Sintang  (south-west  Borneo)  and  10 
females  from  north  Borneo.  He  wrote  that  the  underside  was  bluish  white  speckled  with  pale  brown  in  the 
male  and  thicker  grey-brown  in  the  female.  This  description  does  not  accord  well  with  the  taxon  here 
treated  as  apries.  However,  there  is  a  male  from  Sintang  in  BMNH  labelled  apries  by  Fruhstorfer  and  with 
the  underside  paler  and  greyer  than  usual,  which  presumably  formed  part  of  the  type-series.  In  addition 
there  are  seven  males  of  A.  horsfieldi permagnus  from  Sintang  ex  Fruhstorfer  coll. ,  and  it  seems  likely  that 
these  also  formed  part  of  Fruhstorfer's  type-series  even  though  none  had  been  labelled  by  him  as  apries. 
The  only  other  males  of  apries  ex  coll.  Fruhstorfer  in  BMNH  are  one  labelled  by  Fruhstorfer  as  A.  strigatus 
Moulton  and  one  which  was  apparently  regarded  by  Corbet  as  the  holotype  of  apries  and  is  labelled  /Type 
[red]/Type  [Fruhstorfer  orange]/Kina  Balu  ex  coll.  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/ 
apries/.  As  it  did  not  come  from  Sintang  it  cannot  have  formed  part  of  the  type-series,  and  I  reject  it  as  a 
type.  In  its  place  I  designate  as  lectotype  of  apries  the  male  from  Sintang,  which  is  labelled  /Sintang  Dr 
Martin  H.  Fruhstorfer/21.IV.10/Fruhstorfer  Coll.  B.M.  1933-131/apries  Fr.  [in  Fruhstorfer's  hand]/A. 
apries  apries  Det.  by  Dr.  A.  S.  Corbet  [in  Corbet's  hand]/. 

There  are  also  two  females  of  apries  in  BMNH  which  have  been  treated  as  syntypes.  One  is  labelled 
/Type  [red]/Type  [Fruhstorfer  orange]/Kina  Balu  Borneo/Fruhstorfer  Coll.  B.M.  1933-131/horsfieldi 
apries  Frhst.  [in  Fruhstorfer's  hand]/  and  is  a  paralectotype.  The  other,  labelled  /Type  [red]/Type 
[Fruhstorfer  orange]/Sintang  10.IV.10/Fruhstorfer  Coll.  B.M.  1933-131/  is  rejected  as  a  paralectotype  as  it 
did  not  come  from  north  Borneo  and  therefore  cannot  have  formed  part  of  the  type-series. 


Fig.  27    Allotinus  (Paragerydus)  apries  apries  Fruhstorfer;  Malay  Peninsula.  Male  genitalia.  Lower  left, 
lateral  view  of  phallus. 


44 


J.  N.  ELIOT 


I  designate  as  lectotype  of  eupalion  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
CMB  II.94/Fruhstorfer  Coll.  B.M.  1933-131/strigatus  eupalion  Frhst.  [in  Fruhstorfer's  hand]/GENITA- 
LIA  Slide  No.  ASC  9  Allotinus/. 

Here  it  may  be  mentioned  that  the  female  figured  by  Fruhstorfer  (1916:  pi.  141g)  as  apries  is  a  different 
species,  either  A.  horsfieldi  or  A.  melos,  and  also  that  the  male  and  female  figured  by  Swinhoe  (1910:  pi. 
617,  figs  1,  la)  as  A.  horsfieldi  are  in  BMNH  and  are  in  fact  Bornean  specimens  of  A.  apries  and  A.  melos 
respectively. 

DISTRIBUTION.  Borneo,  including  Pulo  Laut;  Malay  Peninsula;  Sumatra. 

Allotinus  (Paragerydus)  apries  dositheus  Fruhstorfer 

Allotinus  strigatus  dositheus  Fruhstorfer,  1914:  22;  1916:  813.  LECTOTYPE  cT,  JAVA  (BMNH),  here 

designated  [examined]. 

[Allotinus  strigatus  Moulton  sensu  Piepers  &  Snellen,  1918:  22,  pi.  20,  fig.  17  cf  •  Misidentification.] 
Allotinus  apries  dositheus  Fruhstorfer;  Corbet,  19396:  72. 

The  male  does  not  differ  from  that  sex  of  the  nominate  subspecies.  The  female  differs  in  being  brown 
without  a  reddish  tint,  in  this  respect  showing  parallel  variation  with  the  female  of  A.  horsfieldi. 

I  designate  as  lectotype  of  dositheus  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Java  Occident.  Sukabumi  2000'  ex  coll.  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/strigatus  dositheus 
Frhst.  [in  Fruhstorfer's  hand]/GENITALIA  Slide  No.  ASC  8  Allotinus/.  A  female,  similarly  labelled, 
which  has  been  treated  as  a  syntype,  is  a  misidentified  example  of  A.  portunusportunus. 

DISTRIBUTION.  Java. 

Allotinus  (Paragerydus)  apries  ristus  subsp.  n. 

(Fig.  106  cf ) 

Cf  on  the  upperside  differs  from  subsp.  apries  in  having  a  narrower  forewing  visual  brand  1-5  mm  wide.  The 
underside  differs  by  its  greyish  white  ground  colour  and  darker  greyish  brown  markings.  The  striations  are 
fine  and  the  postdiscal  spots  small. 

MATERIAL  EXAMINED 
Holotype  cf ,  Philippines:  central  Palawan,  Languan,  i.1981  (Treadaway  Coll.). 

Allotinus  (Paragerydus)  corbeti  Eliot 

(Fig.  28,  cf  genitalia) 

[Allotinus  dilutus  Corbet  sensu  Corbet,  19406:  pi.  1,  figs  13  cf ,  14  $.  Misidentification.] 
Allotinus  corbeti  Eliot,  1956:  34;  1984:  100.  Holotype  cf ,  WEST  MALAYSIA  (BMNH)  [examined]. 


Fig.  28    Allotinus  (Paragerydus)  corbeti  Eliot;  Borneo:  Pulo  Laut.  Male  genitalia. 


THE  MILETINI  45 

[Allotinus  felderi  felderi  Semper  sensu  Eliot,  1967:  70,  partim.  Misidentification.] 

Allotinus  felderi  corbeti  Eliot;  Eliot,  1967:  70,  fig.  2  cf  genitalia;  1978:  240;  Fleming,  1975:  21,  pi.  21,  fig. 
L41cf. 

This  very  small  species  has  an  uniform  reddish  brown  upperside  in  both  sexes,  and  the  underside  marked  as 
in  A.  leogoron  leogoron.  In  the  male  the  forewing  apex  is  rounded,  and  the  swelling  of  vein  M3  is  confined 
to  the  basal  quarter  of  the  vein;  there  is  no  visual  brand.  The  female  has  the  hindwing  termen  more  strongly 
crenulate  than  A  leogoron. 

Although  smaller,  it  is  possible  that  A  corbeti  might  be  confused  with  A  nicholsi,  since  the  males  of  both 
species  are  similar  on  the  upperside.  Females  are  also  a  similar  shade  of  uniform  reddish  brown,  but  those 
of  corbeti  are  readily  separated  by  their  crenulate  hindwing  with  the  cilia  elongated  into  tufts  at  the  vein 
endings.  In  addition,  corbeti  has  no  humeral  vein  on  the  hindwing,  this  vein  being  present  in  all  six 
examples  of  nicholsi  which  I  have  examined. 

DISTRIBUTION.  West  Malaysia;  Singapore;  Sumatra;  Pulo  Laut;  Mindanao;  Peninsular  Thailand  (Pinrata- 
na).  The  species  appears  to  be  extremely  rare,  but  is  perhaps  often  overlooked. 

Allotinus  (Paragerydus)  unicolorC.  &  R.  Felder 

(Figs  29,  30,  O"  genitalia) 
Allotinus  unicolor  C.  &  R.  Felder,  [1865]:  286. 

This,  the  most  wide-ranging  species  of  the  genus,  shows  great  individual  variation  in  size,  wing  shape  and 
underside  pattern.  In  general  the  smaller  males,  with  forewing  length  as  little  as  11-0  mm,  have  a  more 
rounded  forewing  apex  and  termen  and  a  relatively  shorter  brand,  whilst  the  bigger  males  have  a  more 
acute  apex  and  straighter  termen  and  a  brand  which  is  slightly  longer  and  narrower  in  relation  to  wing  span. 
Examples,  such  as  the  holotype  of  unicolor,  in  which  the  usual  lycaenid  markings  on  the  underside  are 
heavy,  are  comparatively  sparsely  striated.  In  examples  in  which  the  underside  is  more  densely  striated 
there  is  usually  a  reduction  in  the  size  and  darkness  of  these  markings.  On  the  forewing  the  postdiscal  series 
is  typically  on  an  almost  even  curve,  but  examples  in  which  it  is  dislocated  at  vein  M3  to  a  greater  or  less 
extent  are  equally  frequent. 

The  best  characters  for  identifying  the  species  are  the  male  genitalia  (Figs  29, 30)  with  a  broad  valva  with 
the  terminal  process  almost  in  the  centre  line,  the  short  stalk  of  veins  MI  and  R5  of  the  forewing  and  the 
relatively  short  swelling  of  vein  M3  in  the  male. 

Fruhstorfer  grouped  the  complex  into  three  sympatric  'species':  unicolor,  aphocha  andposidion.  Under 
unicolor  he  grouped  comparatively  small  specimens  with  a  whitish  ground  colour  on  the  underside  and 
heavy  postdiscal  markings,  and  under  aphocha  specimens  of  similar  size  and  wing  shape  but  with  less 
pronounced  markings.  He  reserved  posidion  for  large  specimens  in  which  the  male  forewing  was  more 
produced.  Corbet  (1939ft),  though  at  first  inclined  to  agree  with  Fruhstorfer's  arrangement,  finally  decided 
that  all  three  constituted  a  single,  variable  species,  but  at  the  same  time  erected  a  new  'species',  A  dilutus, 


Fig.  29    Allotinus  (Paragerydus)  unicolor  continentalis  Fruhstorfer;  Burma.  Male  genitalia. 


46 


J.  N.  ELIOT 


Fig.  30    Allotinus  (Paragerydus)  unicolor  zitema  Fruhstorfer;  Sulawesi.  Male  genitalia. 

which  he  later  (1956)  relegated  to  subspecies,  for  very  small  examples  from  the  Malay  Peninsula.  The 
various  sizes,  wing  shapes  and  patterns  are  connected iby  intermediates  and,  like  Corbet,  I  can  find  no 
differences  in  the  male  genitalia  of  the  largest  and  smallest  specimens.  Therefore  I  also  consider  that  there 
is  only  one  species,  to  which  I  now  add  the  taxon  continentalis ,  which  has  previously  been  treated  as  a 
subspecies  of  A.  horsfieldi  or  as  a  distinct  species. 

It  is  extraordinarily  difficult  to  decide  what  constitutes  a  valid  subspecies  because  of  the  great  range  of 
phenetic  variation.  When  large  series  are  available  for  comparison,  as  in  coll.  BMNH,  it  is  possible  to 
detect  differences  between  the  average  phenotype  of  different  geographical  areas,  but  because  of  the 
overlap  of  phenetic  characters  it  may  often  be  impossible  to  ascribe  individual  specimens,  if  deprived  of 
their  locality  labels,  to  any  particular  country  of  origin.  I  have  opted  to  retain  as  subspecies  those  groups  of 
populations  which  can  be  distinguished  by  a  mean  of  differences  and  which  are  contained  within  generally 
accepted  faunal  areas  in  preference  to  lumping  together  into  just  two  or  three  polytypic  subspecies  the 
populations  of  widely  separated  geographical  areas  which  in  all  probability  differ  genetically  to  a 
considerable  degree.  In  using  the  key  below  this  limitation  should  be  borne  in  mind. 

The  species  ranges  from  Assam  to  the  Lesser  Sunda  Is. ,  Philippines  and  Sulawesi  (including  the  Sula  Is.) , 
and  is  most  abundant  in  low  level  primary  or  secondary  forest. 

Key  to  the  subspecies  of  A.  (P.)  unicolor 

1  Underside  of  hindwing  with  the  postdiscal  spot  in  space  6  well  inside  that  in  space  7  and  often 

mid-way  to  the  end-cell  spot 2 

Underside  of  hindwing  with  the  postdiscal  spot  in  space  6  much  closer  to  the  spot  in  space  7  than 
to  the  end-cell  spot 7 

2  Underside  ground  colour  greyish  white .  cf  vein  M3  of  fore  wing  swollen  for  not  more  than  half  its 

length.  £  upperside  of  forewing  with  disc  not  conspicuously  paler 3 

Underside  ground  colour  pale  buff,  cf  vein  M3  swollen  for  a  little  over  half  its  length.  $  forewing 
disc  conspicuously  paler  and  often  sullied  whitish unicolor  continentalis  (p.  47) 

3  cf  forewing  brand  1  -0  mm  or  more  wide.  Continental  Asia  and  Borneo 4 

Cf  brand  less  than  1-0  mm  wide.  Philippines  and  Sulawesi 6 

4  9  upperside  reddish  brown 5 

9  upperside  brown  without  a  reddish  tint unicolor  rekkia  (p.  48) 

5  cf  forewing  brand  about  1-0  mm  wide unicolor  unicolor  (p.  47) 

Cf  brand  comparatively  short  and  broad,  about  1-75  mm  wide  in  large  specimens. 

Underside  rather  lightly  marked unicolor  moorei  (p.  48) 

6  Underside  with  postdiscal  markings  usually  heavy  and  sharply  defined,  cf  vein  A/3  swollen  for 

just  under  half  its  length,  brand  sharply  defined unicolor  georgius  (p.  49) 

Underside  with  postdiscal  markings  usually  rather  light  and  tending  to  be  blurred,  cf  vein  M3 
swollen  for  half  its  length  and  brand  usually  rather  inconspicuous unicolor  zitema  (p.  50) 


THE  MILETINI  47 

7    Upperside  hindwing  with  postdiscal  spot  in  space  6  often  below  and  conjoined  to  the  spot  in 

space  7.  $  upperside  brown  without  a  reddish  tint,  forewing  disc  paler . . .      unicolor  postilion  (p.  48) 
Underside  hindwing  with  the  postdiscal  spot  in  space  6  nearly  always  inside  the  spot  in  space  7. 

9  upperside  with  a  slight  reddish  tint,  forewing  disc  only  slightly  paler...       unicolor  aphocha  (p.  48) 

Allotinus  (Paragerydus)  unicolor  unicolor  C.  &  R.  Felder 

Allotinus  unicolor  C.  &  R.  Felder,  [1865]:  286.  Holotype  'cf'  recte  $,  SINGAPORE  (BMNH)  [examined]. 

[Allotinus posidion  myriandus  Fruhstorfer,  1913:  368  (partim);  1916:  811  (partim).] 

Allotinus posidion  eurytanus  Fruhstorfer,  1913:  368;  1916:  811.  LECTOTYPE  cf ,  BORNEO  (BMNH),  here 

designated  [examined].  Syn.  n. 
Allotinus  unicolor  unicolor  Felder;  Fruhstorfer,  1913:  369;  1916:  809,  pi.  141i;  Corbet,  19396;  68,  pi.  1,  figs 

3  $  holotype,  4  cf;  1956:  269,  pi.  44,  fig.  153  cf;  Eliot,  1978:  240,  pi.  20,  figs  5  cf,  6  $. 
[Allotinus  aphocha  aphocha  (Kheil);  Fruhstorfer,  1913:  370  (partim);  1916:  810  (partim).] 
Allotinus  unicolor  eurytanus  Fruhstorfer;  Corbet,  19396:  70,  pi.  1,  figs  9  cf  'holotype  of  eurytanus'  (in 

error),  10  cf  'holotype  of  eurytanus  f.  rebilus'  (in  error). 

Allotinus  dilutus  Corbet,  19396:  70.  Holotype  cf ,  WEST  MALAYSIA  (BMNH)  [examined].  Syn.  n. 
Allotinus  unicolor  dilutus  Corbet;  Corbet,  1956:  269;  Cantlie,  1967:  27;  Fleming,  1975:  21,  pi.  57,  fig. 

L40cf. 

The  female  holotype,  figured  by  Corbet  (19396)  and  mistaken  by  the  Felders  for  a  male,  is  an  atypical 
specimen  with  heavy  markings  and  a  whiter  than  usual  ground  colour.  Such  specimens  occur  most  often  in 
Singapore,  where  they  are  connected  by  intermediates  to  normal  phenotypes.  The  male  figured  by  Corbet 
(19396),  which  matches  the  holotype  fairly  well  but  has  a  slightly  greyer  ground  colour  on  the  underside  (as 
usual  in  males),  would  probably  have  been  identified  by  Fruhstorfer  as  A.  posidion  myriandus  because  of 
its  size  and  wing  shape. 

In  general  Bornean  examples  have  the  lycaenid  markings  smaller  than  in  those  from  continental  Asia, 
but  compensate  by  being  slightly  more  densely  striated.  This  tendency  is  most  extreme  in  examples  from 
south-west  Borneo,  named  eurytanus  by  Fruhstorfer,  which  are  particularly  densely  striated  but  can  be 
matched  by  occasional  examples  from  other  areas. 

I  designate  as  lectotype  of  eurytanus  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Sintang  19.IV.10/Fruhstorfer  Coll.  B.M.  1933-131/posidion  eurytanus  Fr.  [in  Fruhstorfer's  hand]/.  It  was 
figured  by  Corbet  (19396:  fig.  10)  in  error  as  the  holotype  of  A.  unicolor  eurytanus  f.  rebilus.  The  smaller 
specimen  figured  by  Corbet  at  fig.  9  in  error  as  the  holotype  of  eurytanus  is  labelled  /Type  [red]/Type 
[Fruhstorfer  orangeJ/Sintang  26.IV.10/aphocha  rebilus  Frhst.  [in  Fruhstorfer's  hand]/GENITALIA  Slide 
No.  ASC  14  Allotinus/.  Despite  Fruhstorfer's  identification  label  it  cannot  be  accepted  as  lectotype  of 
rebilus,  since  this  taxon  was  described  from  North  Borneo;  it  probably  formed  part  of  Fruhstorfer's  original 
type-series  of  four  males  and  one  female  of  A.  posidion  eurytanus,  and  how  it  obtained  its  label  as  rebilus  is 
a  mystery. 

DISTRIBUTION.  This  very  variable  subspecies  is  found  in  south  Burma,  where  it  has  a  zone  of  intergradation 
with  subsp.  continentalis  between  Rangoon  and  Tavoy,  and  in  Peninsular  Thailand,  West  Malaysia, 
Singapore,  Lingga  Is.,  Natuna  Is.  and  throughout  Borneo,  except  in  the  Kina  Balu  area  where  it  merges 
into  subsp.  moorei. 

Allotinus  (Paragerydus)  unicolor  continentalis  Fruhstorfer 

(Fig.  29,  cf  genitalia) 

[Paragerydus  horsfieldi  (Moore)  sensu  de  Niceville,  1890:  26,  pi.  36,  fig.  156  cf .  Misidentification.] 
[Allotinus  horsfieldi  (Moore)  sensu  Bingham,  1907:  287,  299,  fig.  73  cf ;  sensu  Swinhoe,  1910: 198  (partim, 

nee  pi.  617,  figs  1,  la,  Ib,  Ic).  Misidentifications.] 
Allotinus  horsfieldi  continentalis  Fruhstorfer,  1913:  344;  1916:  812;  Evans,  1932:  212;  Cantlie,  1963:  27. 

Holotype  cf ,  BURMA:  Bhamo  (probably  in  ZSI). 
[Allotinus posidion  subsp.;  Fruhstorfer,  1913:  368.] 
Allotinus  posidion  atacinus  Fruhstorfer,  1916:  811;  Evans,  1932:  212.  Holotype  $.  BURMA  (BMNH) 

[examined].  [Synonymised  by  Eliot,  1967:  70.] 

Allotinus  unicolor  atacinus  Fruhstorfer;  Corbet,  19396:  68,  pi.  1,  fig.  2  $  holotype;  Cantlie,  1963:  27. 
Allotinus  continentalis  continentalis  Fruhstorfer;  Corbet,  19396:  72. 
Allotinus  continentalis  Fruhstorfer;  Eliot,  1967:  70,  fig.  4  cf  genitalia. 


48  J.  N.  ELIOT 

This  is  the  most  distinctive  and  largest  subspecies  of  A.  unicolor,  with  the  forewing  length  of  the  male 
usually  18-19  mm.  On  the  underside  both  sexes  are  pale  buff.  On  the  upperside  the  male  brand  is  longer 
and  wider  than  in  the  other  subspecies,  usually  2-0  mm  wide  and  touching  vein  Cu}  at  its  origin,  while  the 
swelling  of  vein  M3  is  a  little  over  half  its  length.  The  female  is  distinguished  by  the  prominently  paler  discal 
area  on  the  upperside  of  the  forewing,  which  may  be  sullied  whitish  in  the  dry  season. 

DISTRIBUTION.  Assam;  Burma  as  far  south  as  Tavoy;  north-west  Thailand.  Around  the  latitude  of  Rangoon 
it  intergrades  with  subsp.  unicolor. 

Allotinus  (Paragerydus)  unicolor  rekkia  Riley  &  Godfrey 

Allotinus posidion  rekkia  Riley  &  Godfrey,  1921:  180,  pi.  6,  figs  1  d",  2  $.  Holotype  cf ,  THAILAND:  east 

(BMNH)  [examined]. 
Alliotinus  unicolor  rekkia  Riley  &  Godfrey;  Eliot,  1967:  70. 

Females  are  brown  without  the  reddish  tint  of  subsp.  unicolor,  but  otherwise  the  subspecies  does  not  differ 
and  is  of  doubtful  validity. 

DISTRIBUTION.  Only  known  from  eastern  Thailand,  but  probably  also  occurs  in  Cambodia,  Laos  and 
Vietnam. 

Allotinus  (Paragerydus)  unicolor  moorei  (H.  H.  Druce) 

Paragerydus  moorei  H.  H.  Druce,  1895:  562,  pi.  31,  figs  5  cf ,  6  $ .  Syntypes,  BORNEO:  Mt  Kina  Balu  (coll. 

Staudinger,  probably  in  MNHU). 

Allotinus paetus  moorei  (Druce)  Fruhstorfer,  1913:  369: 1916;  811. 
Allotinus  aphocha  rebilus  Fruhstorfer,  1913:  370;  1916:  810.  LECTOTYPE  $,  BORNEO  (BMNH),  here 

designated  [examined].  Syn.  n. 
Allotinus  moorei  (H.  H.  Druce);  Corbet,  19396:  68. 

Druce's  figure  of  the  male  shows  a  comparatively  large  specimen  weakly  marked  beneath  and  with  the 
brand  on  the  upperside  of  the  forewing  rather  short,  broad  and  diffuse.  Judging  by  material  in  BMNH  such 
males  are  rare,  and  are  connected  by  intermediates  to  smaller  males  which  differ  little,  if  at  all,  from 
nominate  unicolor.  The  subspecies  is  therefore  of  doubtful  validity. 

Fruhstorfer's  taxon  rebilus  is  applicable  to  smaller  examples.  I  designate  as  lectotype  a  female  in  BMNH 
labelled  /Type  [red]/Kina  Balu/ex  coll.  H.  Fruhstorfer/ Allotinus  aphocha  rebilus  Fr.  [in  Corbet's  hand]/. 

DISTRIBUTION.  The  subspecies  flies  on  Mt  Kina  Balu,  and  might  conceivably  be  a  local  modification 
occurring  at  higher  elevations  than  normal  examples  referable  to  subsp.  unicolor. 

Allotinus  (Paragerydus)  unicolor  aphocha  Kheil 

Allotinus  aphocha  Kheil,  1884:  28,  pi.  5,  fig.  30  $.  Holotype  $,  NIAS  (probably  in  MNHU). 

Allotinus  posidion  myriandus  Fruhstorfer,  1913:  368;  1916:  811.  Lectotype  cf,  SUMATRA  (BMNH),  by 

designation  (as  holotype)  of  Corbet,  19396:  68  [examined].  Syn.  n. 

Allotinus  aphocha  aphocha  Kheil;  Fruhstorfer,  1913:  370  (partim);  1916:  810  (partim),  pi.  141g  cf  9- 
Allotinus  unicolor  myriandus  Fruhstorfer;  Corbet,  19396:  68,  pi.  1,  figs  5  cf  holotype,  6  9  allotype. 
Allotinus  unicolor  aphocha  Kheil;  Corbet,  19396:  70,  pi.  1,  fig.  11  $. 

Differs  from  subsp.  unicolor  only  in  that  on  the  underside  of  the  hindwing  the  postdiscal  spot  in  space  6  is 
usually  placed  closer  to  the  spot  in  space  7.  Many  specimens  are  inseparable  from  unicolor,  so  the 
subspecies  is  of  doubtful  validity. 

Fruhstorfer  did  not  designate  a  type  of  myriandus,  but  Corbet  (19396)  figured  its  'holotype'  male  and 
'allotype'  female,  and  this  action  constitutes  a  valid  lectotype  selection. 

DISTRIBUTION.  Sumatra;  Bangka  I.;  Batu  Is;  Mentawai  Is;  Nias  I. 

Allotinus  (Paragerydus)  unicolor  posidion  Fruhstorfer 

Allotinus  posidion  posidion  Fruhstorfer,  1913:  368;  1916:  811.  Lectotype  cf,  JAVA:  west  (BMNH),  by 

designation  (as  holotype)  of  Corbet,  19396:  68  [examined]. 
Allotinus  posidion  molionides  Fruhstorfer,  1913:  368;  1916:  811.  Lectotype  cf,  BALI  (BMNH),  by 

designation  (as  holotype)  of  Corbet,  19396:  70  [examined].  Syn.  n. 


THE  MILETINI  49 

Allotinus  posidion  niceratus  Fruhstorfer,  1913:  368;  1916:  812.  LECTOTYPE  cf,  SUMBAWA  (BMNH), 

here  designated  [examined].  Syn.  n. 
Allotinus  unicolor  enganicus  Fruhstorfer,  1913:  369;  1916:  811.  Lectotype  d",  ENGANO  I.  (BMNH),  by 

designation  (as  holotype)  of  Corbet,  19396:  70,  pi.  1,  fig. 12  [examined].  Syn.  n. 
Allotinus  unicolor  bajanus  Fruhstorfer,  1913:  369;  1916:  811;  Corbet,  19396:  70.  LECTOTYPE  cf, 

LOMBOK  (BMNH),  here  designated  [examined].  Syn.  n. 
Allotinus  aphocha  enatheus  Fruhstorfer,  1913:  370;  1916:  810.  LECTOTYPE  $,  JAVA  (BMNH),  here 

designated  [examined].  [Synonymised  by  Corbet,  19396:  70.] 

Allotinus  horsfieldii  [sic]  f.  posidion  Fruhstorfer;  Piepers  &  Snellen,  1918:  12,  pi.  19,  figs  14a  cf ,  14b  $. 
Allotinus suka  Piepers  &  Snellen,  1918: 13  (partim),  pi.  27,  fig.  181  cf .  LECTOTYPE  cf ,  JAVA  (probably  in 

RNH),  here  designated.  [Aberration  reduced  to  infrasubspecific  status  by  Corbet,  19396:  70.] 
Allotinus  unicolor  Felder;  Piepers  &  Snellen,  1918:  14,  pi.  19,  figs  13a  cf ,  13b  ?. 
[Allotinus  aphocha  Kheil;  Piepers  &  Snellen,  1918:  15,  pi.  19,  figs  15a  cf ,  15b  $.] 
Allotinus  unicolor  posidion  Fruhstorfer;  Corbet,  19396:  68,  pi.  1,  figs  7  cf  holotype,  8  $  'allotype'  of 

enatheus. 

Allotinus  unicolor  molionides  Fruhstorfer;  Corbet,  19396:  70,  pi.  1,  fig.  13  cf  holotype. 
Allotinus  unicolor  bajanus  Fruhstorfer;  Corbet,  19396:  70. 

The  female  lacks  the  reddish  tint  of  subsp.  myriandus  and  usually  the  forewing  disc  is  paler.  On  the 
underside  of  the  hindwing  the  postdiscal  spot  in  space  6  is  more  often  directly  below,  and  conjoined  to,  the 
spot  in  space  7;  this  character  occurs  most  often  in  examples  from  the  Lesser  Sunda  Is. ,  which  also  have  the 
postdiscal  spots  on  average  heavier  than  in  Javanese  examples. 

Fruhstorfer  did  not  designate  types  of  his  taxa,  but  Corbet  (19396)  figured  the  'holotype'  males  of 
posidion,  molionides  and  enganicus,  and  this  action  constitutes  valid  lectotype  selections.  He  also  figured  a 
female  which  he  described  as  A.  unicolor  posidion  f.  enatheus  Fruh.  $  allotype.  I  now  designate  this  female 
as  lectotype  of  enatheus;  it  is  labelled:  /Type  [red]/Type  [Fruhstorfer  orange]/Java  Occident.  Sukabumi 
2000'  ex  coll.  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/aphocha  enatheus  Fr.  [in  Fruhstorfer's 
hand]/. 

I  designate  as  lectotype  of  niceratus  a  male  in  BMNH  labelled  /Type  [red]/Sumbawa/Allotinus  posidion 
niceratus  Fruh.  Type  [in  Corbet's  hand]/Adams  Bequest  B.M.  1912-399/.  It  is  to  be  presumed  that 
Fruhstorfer  saw  this  specimen  during  his  visit  to  BMNH  prior  to  publication  of  his  1913  paper,  as  there  are 
no  specimens  from  Sumbawa  from  his  collection  in  BMNH. 

I  designate  as  lectotype  of  bajanus  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Pringabaja  April  1896  H.  Fruhstorfer/unicolor  bajanus  Frhst.  [in  Fruhstorfer's  hand]/. 

Piepers  &  Snellen  (1918)  figured  two  different  species  as  Allotinus  suka:  a  female  on  pi.  19,  fig.  12b  to 
which  they  had  previously  referred  as  Allotinus  horsfieldii  and  a  male  on  pi.  27,  fig.  181.  When  relegating 
suka  to  the  status  of  a  form  of  A.  unicolor  posidion,  in  which  the  usual  striations  of  the  underside  are 
absent,  Corbet  (19396:  70)  referred  only  to  the  latter  figure  which  he  said,  in  error,  represented  a  female. 
Corbet's  action  could,  perhaps,  be  taken  to  restrict  the  name  suka  to  the  species  represented  in  fig.  181 ,  but 
to  make  absolutely  certain  that  there  should  be  no  confusion  in  the  application  of  the  name  I  now  designate 
this  male  as  lectotype.  It  should  be  in  RNH. 

DISTRIBUTION.  Java;  Bali;  Lombok;  Sumbawa;  Engano  I. 

Allotinus  (Paragerydus)  unicolor  georgius  Fruhstorfer 

[Paragerydus  horsfieldi  (Moore)  sensu  Semper,  1889: 164,  partim.  pi.  31,  fig.  18  cf  •] 

Allotinus  posidion  georgius  Fruhstorfer,  1913:  368,  partim;  1916:  812.  Holotype  cf,  PHILIPPINES:  Bohol 

(SM)  [examined]. 
Allotinus  unicolor  leitus  Fruhstorfer,  1916:  811.  Holotype  $,  PHILIPPINES:  Mindoro  (coll.  Staudinger, 

probably  in  MNHU). 

Fruhstorfer  named  georgius  from  Semper's  records  and  figures  of  'horsfieldi'  from  Bohol  and  Mindanao, 
which  he  subsequently  (1916)  realised  represented  two  species.  Thereupon  he  restricted  georgius  to  Bohol, 
so  that  this  name  applies  to  the  male  figured  by  Semper  at  fig.  18,  which  automatically  becomes  the 
holotype;  it  is  labelled  /18/213/Coll.  C.  Semper/reverdini/original  of  Semper  PI.  31,  fig.  18  cf  from  Bohol. 
Holotype  of  Allotinus  posidion  georgius  Fruh.  det.  J.  N.  Eliot  ix.1982/.  It  is  heavily  marked  below  much  as 
in  the  holotype  of  unicolor,  but  on  the  upperside  the  brand  is  a  little  shorter  and  narrower  than  in  that 
subspecies. 
Fruhstorfer  named  leitus  from  a  single  female  with  a  yellowish  discal  area  on  the  forewing  and  heavy 


50 


J.  N.  ELIOT 


markings  on  the  underside.  The  latter  character  suggests  that  it  pertains  to  the  same  subspecies  as  georgius, 
but  as  I  have  seen  no  examples  from  Mindoro  it  is  provisionally  placed  in  synonymy. 

DISTRIBUTION.  A  unicolor  must  be  very  rare  in  the  Philippines,  as  it  is  not  represented  in  coll.  Treadaway, 
and  in  BMNH  there  are  only  two  females,  one  of  which  is  labelled  'Philippines  Pryer'.  This  appears  to  be  an 
aberration;  it  is  lightly  striated  on  the  underside,  as  in  examples  from  north  Borneo,  and  the  postdiscal 
markings  are  elongated  into  longitudinal  streaks.  It  may  have  come  from  the  Sulu  Is. ,  which  were  visited  by 
Pryer,  or  may  be  a  mislabelled  example  from  Borneo,  where  Pryer  did  nearly  all  his  collecting. 

Allotinus  (Paragerydus)  unicolor  zitema  Fruhstorfer 
(Fig.  30,  cf  genitalia) 

Allotinus  aphocha  zitema  Fruhstorfer,  1916:  810.  LECTOTYPE  $,  SULAWESI  (BMNH),  here  designated 
[examined]. 

The  subspecies  is  distinguished  by  the  male  brand,  which  is  a  little  narrower  and  more  obscure  than  in  other 
subspecies. 

I  designate  as  lectotype  a  female  in  BMNH  labelled:  Type  [red]/Type  [Fruhstorfer  orange]/Nord- 
Celebes  Toli  Toli  Nov.  -  Dez.  1895  H.  Fruhstorfer/aphocha  zitema  Fr.  [in  Fruhstorfer's  hand]/Fruhstorfer 
Coll.  B.M.  1933-131/.  I  have  seen  no  males  ex  Fruhstorfer  coll. 

DISTRIBUTION.  Sulawesi  and  the  Sula  Is. 

Allotinus  (Paragerydus)  paetus  (de  Niceville) 
(Fig.  31  cf  genitalia) 

Paragerydus  paetus  de  Niceville,  1895:  269,  pi.  O,  fig.  12  cf .  Syntypes,  SUMATRA:  north-east  (probably  in 

ZSI). 

Allotinus  paetus  paetus  (de  Niceville)  Fruhstorfer,  1913:  369;  1916:  811,  pi.  141i  <j>. 
Allotinus  paetus  (de  Niceville);  Corbet,  19396:  68,  pi.  1,  fig.  1  cf . 

This  species  bears  a  fairly  close  resemblance  to  examples  of  A.  unicolor  which  are  strongly  marked 
beneath,  as  in  the  holotype  of  unicolor.  But  it  is  larger,  with  forewing  length  averaging  18-19  mm  in  males, 
the  forewing  brand  and  swelling  of  vein  A/3  are  slightly  longer  than  in  all  unicolor  subspecies  apart  from 
subsp.  continental,  being  just  over  half  the  length  of  the  vein,  and  on  the  underside  the  ground  colour  is 
more  chalky  whitish.  The  male  genitalia  are  rather  similar  to  those  of  A.  unicolor,  but  the  phallus  is  stouter 
and  the  distal  portion  of  the  valva  narrower,  with  the  terminal  process  not  curved  up  so  strongly  towards 
the  centre  line. 

DISTRIBUTION.  The  species  is  only  known  from  Sumatra,  where  it  appears  to  fly  in  the  Barisan  Range  from 
the  Battak  mountains  in  the  north  to  the  extreme  south,  where  it  was  taken  in  numbers  by  Doherty. 


Fig.  31     Allotinus  (Paragerydus) paetus  (de  Niceville);  Sumatra.  Male  genitalia. 


THE  MILETINI 


51 


Fig.  32    Allotinus  (Paragerydus)  parapus  Fruhstorfer;  Borneo.  Male  genitalia.  Lower  right,  ventral  view 
of  valvae  and  phallus. 

Allotinus  (Paragerydus)  parapus  Fruhstorfer 

(Fig.  32,  cf  genitalia) 

Allotinus  parapus  Fruhstorfer,  1913:  343;  1916:  809,  pi.  141h  cf;  Corbet,  19396:  66,  fig.  8  cf  genitalia. 
LECTOTYPE  cf ,  BORNEO  (BMNH),  here  designated  [examined]. 

The  sexes  are  alike  in  wing  shape  and  in  having  a  rounded  hindwing  termen  with  the  cilia  inconspicuously 
elongated  at  the  vein  endings.  In  the  male  vein  M3  is  swollen  for  just  under  half  its  length  and  clothed  with 
the  usual  specialised  scales.  The  white  forewing  b<  >rdered  with  dark  brown  and  the  paler  brown  hindwing 
render  the  species  unmistakable. 

Both  Fruhstorfer  and  Corbet,  whose  figure  of  the  male  valva  is  completely  misleading,  stated  that  vein 
M3  was  not  swollen  in  the  male,  and  on  this  account  placed  the  species  in  the /a//ajt-group  (Artengruppe 
Allotinus).  In  fact  the  swelling  is  as  well  developed  as  in  some  other  species  of  the  subgenus,  but  because 
the  surrounding  area  is  white  no  visual  brand  is  apparent. 

I  designate  as  lectotype  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/Fruhstorfer 
Coll.  B.M.  1933-131/Nord-Borneo/ Allotinus  parapus  Frhst.  [in  Fruhstorfer's  hand]/.  A  female,  similarly 
labelled,  is  a  paralectotype. 

DISTRIBUTION.  The  species  is  montane  and  is  known  only  from  Mt  Kinabalu  in  Sabah. 

Allotinus  (Paragerydus)  nivalis  (H.  Druce) 

(Figs  33,  34,  cf  genitalia;  75  cf ) 
Miletus  nivalis  H.  Druce,  1873:  348. 

This  and  the  next  two  taxa  form  a  natural  group  of  small  species  distinguished  by  the  possession  of  a  white 
fleck  at  the  end  of  vein  R5  at  the  apex  of  the  forewing  in  addition  to  the  usual  white  flecks  at  the  ends  of  veins 
Sc,  RI,  R2  and  R3.  In  males  the  swelling  of  vein  M3  is  weak  and  short,  and  there  is  no  visual  brand. 

In  A.  nivalis  the  forewing  termen  is  almost  regular  in  both  sexes,  and  on  the  underside  of  the  hindwing 
the  central  spot  in  space  7  is  not,  or  only  very  little  darkened.  On  average  it  is  larger  than  the  allied  A. 
substrigosus  and  A.  davidis. 

The  species  is  confined  to  Borneo  and  the  Philippines. 

Key  to  the  subspecies  of  A.  (P.)  nivalis 

1    Underside  of  forewing  with  a  more  or  less  developed  submarginal  brownish  blotch  about  1  -0-1  -5 
mm  wide  astride  vein  M3;  postdiscal  series  strongly  dislocated  at  veins  M3  and  Cu2 

nivalis  nivalis  (p.  52) 


52 


J.  N.  ELIOT 


Fig.  33    Allotinus  (Paragerydus)  nivalis  nivalis  (H.  Druce);  Borneo.  Male  genitalia. 

-    Underside  without  a  brownish  blotch  astride  vein  M3;  postdiscal  series  placed  nearer  the  termen 

and  only  a  little  dislocated  at  veins  A/3  and  Cu2 nivalis  felderi  (p.  52) 

Allotinus  (Paragerydus)  nivalis  nivalis  (H.  Druce) 

(Fig.  33,  cf  genitalia) 

Miletus  nivalis  H.  Druce,  1873:  348,  'cf'  recte  <j>.  Holotype  $,  BORNEO  (BMNH)  [examined]. 
Allotinus  nivalis  nivalis  (Druce)  Fruhstorfer,  1913:  370;  1916:  810,  pi.  141g. 
Allotinus  nivalis  (H.  Druce);  Eliot,  1967:  71. 

The  characters  of  the  subspecies  are  given  in  the  key.  The  figure  by  Fruhstorfer  (1916:  141g)  is  very  poor 
and  shows  an  undersized  specimen. 

DISTRIBUTION.  Throughout  Borneo,  including  Pulo  Laut. 

Allotinus  (Paragerydus)  nivalis  felderi  Semper 

(Fig.  34  cf  genitalia;  75  cf ) 

Allotinus  felderi  Semper,  1889: 163,  pi.  31,  fig.  22  $ ;  Fruhstorfer,  1913:  370;  1916:  810.  LECTOTYPE  cf , 
PHILIPPINES  (SM),  here  designated  [examined]. 


Fig.  34    Allotinus  (Paragerydus)  nivalis  felderi  Semper;  Mindanao.  Male  genitalia.  Lower  left,  ventral 
view  of  valvae  and  phallus  of  another  specimen. 


THE  MILETINI  53 

Allotinus  felderi felderi  Semper;  Eliot,  1967:  70,  partim. 
Allotinus  niv  alls  felderi  Semper;  Eliot,  1984:  100. 

The  differences  from  the  nominate  subspecies  are  given  in  the  key. 

Semper  described  the  taxon  from  two  males  from  Mindanao  and  two  females  from  Luzon.  I  designate  as 
lectotype  one  of  the  males  in  SM,  labelled  /Coll.  C.  Semper/Ost  Mind./212/All.  felderi  typ.  Semper/. 

Eliot  (1967)  confused  felderi  with  A.  corbeti  from  a  pair  oicorbeti  and  one  exceptionally  small  female  of 
true  felderi  from  Mindanao,  all  of  which  had  been  placed  under  the  latter  name  in  coll.  BMNH. 

DISTRIBUTION.  Throughout  the  Philippines.  In  addition  to  examples  from  Luzon,  Sibuyan  and  Mindanao  in 
BMNH  I  have  examined  examples  in  coll.  Treadaway  from  Marinduque,  Negros  and  Samar. 

Allotinus  (Paragerydus)  substrigosus  (Moore) 

(Figs  35,  cf  genitalia;  76,  77  cT) 
Logania  substrigosa  Moore,  1884:  22. 

Until  separated  by  Eliot  (1967)  all  authors  treated  this  species  as,  or  as  a  subspecies  of,  A.  nivalis,  from 
which  it  differs  as  follows.  On  average  it  is  smaller,  with  forewing  length  occasionally  as  little  as  9-0  mm.  In 
both  sexes  the  forewing  termen  is  distinctly  crenulate,  becoming  more  exaggerated  in  the  dry  season  form 
from  Burma.  On  the  underside  of  the  hindwing  the  central  spot  in  space  7  is  at  least  partially  blackened  and 
there  is  often  a  black  subbasal  spot  in  space  Ib,  the  corresponding  spots  in  nivalis  not  being  blackened.  The 
male  genitalia  are  very  similar,  but  in  substrigosus  the  phallus  is  narrower  than  in  sympatric  nivalis. 

The  species  ranges  from  central  Burma  and  Thailand  to  Sundaland  and  Mindanao.  In  Borneo  it  appears 
to  be  rarer  than  A.  nivalis. 

Key  to  the  subspecies  of  A.  (P.)  substrigosus 

1  Underside  of  hindwing  with  the  spot  mid-space  7  blackened  only  in  upper  half  of  space,  and  not 

more  than  1  -0  mm  wide 2 

-  Underside  of  hindwing  with  the  spot  mid-space  7  solidly  black,  nearly  2-0  mm  wide  and 

extending  right  across  space substrigosus  ballantinei  (p.  55) 

2  Underside  greyish- white.  Forewing  crenulate 3 

-  Underside  pale  buff.  Forewing  only  very  weakly  crenulate substrigosus  yusukei  (p.  55) 

3  On  underside  of  forewing  the  white  fleck  at  end  of  vein  7  barely  enters  space  6 4 

-  On  underside  of  forewing  the  white  fleck  at  the  end  of  vein  R5  is  continued  as  an  oblique  white 

streak  half-way  across  space  6 substrigosus  substrigosus  (p.  53) 

4  cf  with  vein  M3  swollen  for  only  one-quarter  of  its  length substrigosus  lenaia  (p .  54) 

-  cf  with  vein  M3  swollen  for  one-third  of  its  length,  as  in  subsp.  substrigosus 

substrigosus  sibyllinus  (p.  54) 

Allotinus  (Paragerydus)  substrigosus  substrigosus  (Moore) 

(Fig.  35,  cf  genitalia) 

Logania  substrigosa  Moore,  1884:  22;  1886;  39,  pi.  3,  fig.  8  $.  Holotype  $,  BURMA:  Mergui  Archipelago 

(probably  in  ZSI). 

[Paragerydus  nivalis  (H.  Druce)  sensu  Distant,  1884:  207,  pi.  22,  fig.  11  $.  Misidentification.] 
[Allotinus  nivalis  (H.  Druce)  sensu  de  Niceville,  1890:  30  partim,  pi.  36,  fig.  159  $  holotype  of  substrigosa; 

sensu  Bingham,  1907:  301;  sensu  Swinhoe,  1910:  197,  pi.  616,  figs  2,  2b  cf,  2a  $;  sensu  Piepers  & 

Snellen,  1918: 16,  pi.  20,  fig.  19  cf .  Misidentifications.] 
Allotinus  nivalis  magaris  Fruhstorfer,  1913:  370;  1916:  810.  LECTOTYPE  cf ,  SUMATRA  (BMNH),  here 

designated  [examined].  [Synonymised  by  Eliot,  1967:  71.] 
Allotinus  nivalis  substrigosa  (Moore)  Fruhstorfer,  1916:  810;  Evans,  1932: 212;  Corbet,  19396:  68;  Cantlie, 

1963:  28. 

Allotinus  substrigosa  substrigosa  (Moore);  Eliot,  1967:  71;  1978:  240. 
Allotinus  substrigosus  substrigosus  (Moore);  Fleming,  1975:  21,  pi.  57,  fig.  L38  $  • 

In  the  male  the  swelling  of  vein  M3  extends  to  one-third  of  its  length.  On  the  underside  there  is  a  prominent 
white  streak  at  the  forewing  apex,  and  on  the  hindwing  the  central  spot  in  space  7  is  strongly  blackened. 
From  central  Burma  as  far  south  as  Tavoy  a  distinct  dry  season  form  occurs.  The  forewing  termen  is  more 


54 


J.  N.  ELIOT 


Fig.  35    Allotinus  (Paragerydus)  substrigosus  yusukeisubsp.  n.;  Mindanao.  Male  genitalia.  Lower  left,  A. 
substrigosus  substrigosus  (Moore),  Malay  Peninsula;  right  valva  and  phallus. 

strongly  crenulate;  on  the  underside  the  ground  colour  becomes  more  greyish;  on  the  forewing  the  apex 
and  margin  to  a  depth  of  2-0-3-0  mm  are  shaded  with  brown,  so  that  the  oblique  apical  white  streak  stands 
out  more  conspicuously;  and  on  the  hindwing  there  is  a  similar  brown  area  in  spaces  3,  4  and  5,  and  the 
outer  postdiscal  spot  in  space  7,  and  sometimes  that  in  space  6  also,  may  be  blackened  on  their  inner  edges  - 
a  feature  otherwise  only  found  in  A.  davidis.  This  dry  form,  with  its  mottled  appearance  and  crenulate 
wings,  gives  the  impression  of  a  Logania  species. 

I  designate  as  lectotype  of  magaris  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Selesseh  15.VII.94/Fruhstorfer  Coll.  B.M.  1933-131/Sumatra  Monies  Battak  ex  coll.  Fruhstorfer/nivalis 
magaris  Frhst.  [in  Fruhstorfer's  hand]/.  The  locality  Selesseh  lies  a  little  above  sea  level  in  north-east 
Sumatra  and  is  more  likely  to  be  the  correct  locality  than  the  Battak  Mts. 

DISTRIBUTION.  Burma,  as  far  north  as  east  Pegu;  Thailand;  West  Malaysia;  Sumatra;  Borneo;  Java  (Piepers 
&  Snellen,  1918). 

Allotinus  (Paragerydus)  substrigosus  lenaia  Fruhstorfer 

Allotinus  nivalis  lenaia  Fruhstorfer,  1913:  370;  1916:  810.  LECTOTYPE  cf,  NIAS  (BMNH),  here 

designated  [examined]. 
Allotinus  substrigosa  lenaia  Fruhstorfer;  Eliot,  1967:  72. 

In  the  single  male  which  I  have  seen  the  swelling  of  vein  M3  is  confined  to  the  basal  quarter  of  the  vein.  On 
the  underside  of  the  forewing  the  white  streak  at  the  end  of  vein  R5  at  most  barely  enters  space  6,  and  in 
females  the  markings  are  on  average  lighter  than  in  the  nominate  subspecies. 

I  designate  as  lectotype  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/Nias  ex  coll. 
Fruhstorfer/nivalis  lenaia  Frhst.  [in  Fruhstorfer's  hand]/. 

DISTRIBUTION.  Nias  I. 

Allotinus  (Paragerydus)  substrigosus  sibyllinus  Riley 

Allotinus  nivalis  sibyllina  Riley,  1944:  254,  pi.  2,  fig.  27  cf .  Holotype  cf ,  MENTAWAI  Is.:  Sipora  (BMNH) 
[examined]. 

The  upperside  agrees  with  the  nominate  subspecies.  On  the  underside  the  white  fleck  at  the  forewing  apex 
is  as  redu'ced  as  in  subsp.  lenaia.  On  the  hindwing  the  central  spot  in  space  7  may  be  heavily  blackened,  as  in 
the  nominate  subspecies,  or  only  lightly  blackened.  In  his  original  description  Riley  wrote  that  this  spot  is 
not  blackened,  but  in  seven  out  of  the  eight  specimens  in  the  type-series  it  is  blackened  to  a  greater  or  less 
degree.  The  eighth  specimen,  a  worn  male,  is  so  rubbed  that  it  is  possible  some  black  scales  were  originally 
present. 
The  subspecies  seems  to  be  only  doubtfully  separable  from  subsp.  lenaia. 

DISTRIBUTION.  Mentawai  Is. 


THE  MILETINI  55 

Allotinus  (Paragerydus)  substrigosus  ballantinei  subsp.  n. 

(Fig.  76  cf ) 

Cf  forewing  13-5  mm.  Wing  shape,  as  in  subsp.  substrigosus,  with  forewing  termen  crenulate  and  apex 
rather  pointed.  Swelling  of  vein  M3  inconspicuous,  extending  a  little  under  one-third  of  its  length. 
Underside  very  pale  buff,  rather  densely  striated  but  with  postdiscal  spots  small  or  evanescent,  as  in  some 
individuals  of  subsp.  substrigosus.  Forewing  with  white  speck  at  end  of  vein  R5  not  extending  into  space  6. 
Hindwing  with  spot  in  mid-space  7  exceptionally  large,  rather  quadrate,  nearly  2-0  mm  wide  and  extending 
right  across  space. 

MATERIAL  EXAMINED 
Holotype  cf ,  Philippines:  Palawan,  Port  Barton,  ix.1983  (A  Ballantine)  (BMNH). 

The  subspecies  is  named  after  Dr  Alistair  Ballantine,  who  caught  the  unique  holotype  and  presented  it  to 
BMNH. 

Allotinus  (Paragerydus)  substrigosus  yusukei  subsp.  n. 

(Figs  35  cf  genitalia;  77  cf ) 

Cf  forewing  11-5  mm.  Termen  more  rounded  and  apex  less  pointed  than  in  foregoing  subspecies.  Swelling 
of  vein  M3  short,  extending  over  one-quarter  of  vein,  overlying  specialised  scales  pale  so  that  it  stands  out 
prominently.  Underside  ground  colour  pale  buff;  forewing  with  white  fleck  at  end  of  vein  R5  extending  as 
an  oblique  white  streak  half-way  across  space  6.  Hindwing  with  central  black  spot  in  space  7  narrow  but 
well-defined;  a  small  subbasal  black  spot  in  space  la. 

MATERIAL  EXAMINED 
Holotype  cf ,  Philippines:  Mindanao,  Tandag,  Surigao,  xii.1982  (ex  Takanami  coll.)  (BMNH). 

The  subspecies  is  named  after  Mr  Yusuke  Takanami,  who  generously  presented  the  holotype  and  many 
other  rare  Philippine  lycaenids  to  BMNH. 

Allotinus  (Paragerydus)  davidis  Eliot 
(Fig.  36,  cf  genitalia) 

Allotinus  davidis  Eliot,  1959:  377,  pi.  10,  figs  6  cf ,  7  $ ,  text-fig,  cf  valva;  1978:  240;  Fleming,  1975:  21,  pi. 
57,  fig.  L39  $.  Holotype  cf ,  SINGAPORE  (BMNH)  [examined]. 

The  wing  shape  is  that  of  A.  nivalis,  with  the  forewing  termen  barely  perceptibly  crenulate.  On  the 
upperside  both  sexes  are  reddish  brown.  In  the  male  vein  M3  of  the  forewing  is  swollen  only  in  its  basal 
quarter.  On  the  underside  the  ground  colour  is  pale  buff;  on  the  forewing  the  white  fleck  at  the  apex  does 
not  cross  vein  R5;  on  the  hindwing  the  central  spot  in  space  7  is  blackened  and  the  postdiscal  spot  is  also 
narrowly  blackened  on  its  inner  edge. 

In  the  male  genitalia  the  valva  differs  from  all  others  in  the  subgenus  by  not  having  the  costa  truncate 
before  the  terminal  process,  which  curves  upwards  in  the  manner  of  subgenus  Allotinus.  In  all  other 


Fig.  36    Allotinus  (Paragerydus)  davidis  Eliot;  Singapore.  Male  genitalia. 


56  J.  N.  ELIOT 

respects  the  genitalia  are  typical  of  Paragerydus,  and  the  absence  of  a  humeral  vein  on  the  hindwing  helps 
to  confirm  its  position  in  this  subgenus. 

DISTRIBUTION.  West  Malaysia;  Singapore;  Sumatra  (coll.  Diehl);  Peninsular  Thailand  (Pinratana).  The 
species  appears  to  be  extremely  rare,  but  is  perhaps  sometimes  overlooked  because  of  its  small  size  and 
resemblance  to  A.  substrigosus . 

Allotinus  (Paragerydus)  drumila  (Moore) 

(Fig.  37,  cf  genitalia) 
Miletus  drumila  Moore,  [1866]:  777. 

This  large  species  stands  rather  far  apart  from  the  remainder  of  the  subgenus  in  its  markings  and  in  its 
pronounced  seasonal  variation.  On  the  other  hand  the  male  genitalia,  presence  on  the  forewing  of  a 
swollen  vein  M3  and  absence  of  a  humeral  vein  on  the  hindwing,  are  characteristic  of  Paragerydus,  so  that  it 
seems  unnecessary  to  retain  for  it  the  separate  subgenus  Miletographa. 

On  the  underside  the  appearance  is  of  a  Miletus  species,  with  the  catenulate  markings  characteristic  of 
that  genus;  the  difference  is  that  in  A.  drumila  the  ground  between  the  usual  lycaenid  markings  is  speckled 
with  small,  irregular  pale-edged  spots.  The  upperside  and  especially  the  seasonally  dimorphic  wing  shape 
recall  the  smaller,  sympatric  Miletus  chinensis.  The  wet  season  form  is  brown,  with  a  similar  curved  series 
of  postdiscal  spots  on  the  forewing,  paler  brown  in  the  male,  larger  and  sullied  white  in  the  female.  In  the 
dry  season  the  forewing  acquires  a  sharp  point  at  the  apex  and  prominent  lobe  at  the  tornus,  and  the  mainly 
white  female  is  broadly  similar  to  the  dry  season  M.  chinensis  longeana.  The  main  point  of  dissimilarity  is 
that  in  A.  drumila  the  most  extreme  dry  form  occurs  in  the  Himalayas  and  Assam,  whereas  in  M.  chinensis 
the  most  extreme  dry  form  occurs  in  Burma. 

The  species  is  Indo-Burmese,  submontane  and  apparently  not  rare  where  it  occurs,  but  with  a  restricted 
distribution.  There  are  two  rather  doubtfully  valid  subspecies. 

Key  to  the  subspecies  of  A.  (P.)  drumila 

1     $  dry  season  form  white  with  black  costal  and  marginal  border  on  the  forewing  and  blackish 

costal  area  on  the  hindwing drumila  drumila  (p.  56) 

$  dry  season  form  with  the  white  areas  sullied  with  buff  scales drumila  aphthonius  (p.  57) 

Allotinus  (Paragerydus)  drumila  drumila  (Moore) 

(Fig.  37,  cf  genitalia) 

Miletus  drumila  Moore,  [1866]:  777,  pi.  41,  fig.  12  $.  Holotype  'cf '  recte  <j>,  INDIA:  Sikkim  (not  located). 

[Dry  season  form.] 
Gerydus  drumila  (Moore)  Moore,  1883:  521.  [Wet  season  form.] 


Fig.  37    Allotinus  (Paragerydus)  drumila  drumila  (Moore)  wet  season  form  multistrigatus  de  Niceville; 
Sikkim.  Male  genitalia. 


THE  MILETINI  57 

Allotinus  multistrigatus  de  Niceville,  1886:  253,  pi.  11,  figs  11  o" ,  2  $ ;  de  Niceville,  1890:  29,  pi.  26,  figs  157 
Cf,  158  $;  Bingham,  1907:  298;  Swinhoe,  1910:  195,  pi.  615,  figs  2,  2b  cf ,  2a,  2c  $ ;  Fruhstorfer,  1913: 
371;  1916:  815,  pi.  141h  cf  $.  LECTOTYPE  cf ,  INDIA:  Sikkim  (BMNH),  here  designated  [examined] 
[Wet  season  form.]  [Synonymised  by  Cantlie,  1963:  26.] 

Miletus  insignis  Staudinger,  1888:  269,  pi.  94  'cf'  recte  $.  Holotype  $,  INDIA:  Sikkim  (probably  MNHU). 
[Synonymised  by  de  Niceville,  1890:  28.] 

Allotinus  drumila  (Moore)  de  Niceville,  1890:  28;  Bingham,  1907:  297;  Swinhoe,  1910:  194,  pi.  615,  figs  1, 
Ib  Cf,  la,  Ic  $;  Fruhstorfer,  1913:  371;  1916:  815,  pi.  141i  $;  Evans,  1932:  211. 

Miletographa  drumila  (Moore)  Rober,  1892:  277. 

Allotinus  multistrigatus  multistrigatus  de  Niceville;  Evans,  1932:  242. 

Allotinus  drumila  drumila  (Moore)  with  wet  season  form  multistrigatus  de  Niceville;  Cantlie,  1963:  26,  pi. 
26,  fig.  H. 6. 2. 

Moore  originally  described  the  dry  season  female  in  error  as  the  male.  Later  (1883)  he  realised  his  mistake 
and  correctly  described  the  male  from  the  wet  season  form,  which  de  Niceville  later  redescribed  as  a 
separate  species,  A.  multistrigatus. 

Judging  by  material  in  BMNH,  intermediate  forms,  with  the  wing  shape  of  the  dry  form  drumila,  fly  in 
the  autumn  and  winter,  while  drumila,  with  its  mostly  white  female,  is  really  a  spring  form  flying  from 
February  to  May  (one  female  ex  Fruhstorfer  coll.  is  labelled  June,  but  I  feel  sure  this  is  an  error,  as  the 
type-series  of  multistrigatus  was  caught  in  this  month). 

Moore's  type  of  drumila  cannot  be  traced,  but  there  are  two  wet  season  males  in  BMNH  from  Darjiling 
and  the  Khasi  Hills  from  which  Moore  described  the  male,  both  of  which  bear  BMNH  type-labels.  But  as 
they  did  not  form  part  of  the  original  type-series  they  cannot  be  accepted  as  types. 

The  figures  accompanying  de  Niceville's  original  description  of  multistrigatus  were  taken  from  examples 
in  coll.  Moller,  and  I  designate  as  lectotype  a  male  in  BMNH,  presumably  from  the  original  type-series, 
labelled  /Type  [red]/Sikkim  Moller/ Allotinus  multistrigatus  de  Niceville  cf  TYPE/Rothschild  Bequest 
B.M.  1939-1/. 

DISTRIBUTION.  Kumaon,  in  the  central  Himalayas,  to  Assam. 

Allotinus  (Paragerydus)  drumila  aphthonius  Fruhstorfer 

Allotinus  aphthonius  Fruhstorfer,  1913:  371;  1916:  815.  LECTOTYPE  cf,  BURMA  (BMNH),  here 

designated  [examined].  [Intermediate  form.] 
Allotinus  drumila  grisea  Riley  &  Godfrey,  1921:  180.  Holotype  $,  THAILAND  (BMNH)  [examined].  [Dry 

season  form.]  [Synonymised  by  Pinratana,  1981:  31.] 
Allotinus  multistrigatus  apthonius  [sic]  Fruhstorfer;  Evans,  1932:  242 
Allotinus  drumila  apthonius  [sic]  Fruhstorfer;  Cantlie,  1963:  26;  Pinratana,  1981:  31,  pi.  4,  fig.  19  cf ,  pi.  5, 

figs  10  Cf,  11  $. 

The  subspecies  is  doubtfully  valid,  differing  only  in  that  the  most  extreme  dry  form  of  female  has  the  white 
areas  always  sullied  with  buff  scales,  as  in  the  holotype  of  grisea  and  as  in  Pinratana's  figure  of  the  female. 
The  wet  season  and  intermediate  forms  do  not  differ  from  the  corresponding  forms  of  subsp.  drumila. 

In  BMNH  there  are  one  male  and  two  females  labelled  as  types  of  aphthonius.  I  designate  as  lectotype 
the  male  labelled  /Type  [red]/Type  [Fruhstorfer  orangeJ/Tenasserim  Tandong  4000'  Mai  Fruhstorfer 
leg. /Fruhstorfer  Coll.  B.M.  1933-1/Allotinus  aphthonius  Frhst.  [in  Fruhstorfer's  hand]/.  Two  females, 
similarly  labelled,  except  that  one  bears  a  final  label  /Allotinus  aphthonius  Fruh.  [in  Corbet's  hand]/  in 
place  of  the  label  in  Fruhstorfer's  hand,  are  paralectotypes.  The  male  has  a  pointed  forewing  and  rather 
dentate  hindwing,  and  is  marked  as  in  the  wet  season  form.  Fruhstorfer  stated  that  it  was  a  dry  season  form, 
but  in  fact  it  is  an  intermediate  form  nearer  to  the  wet  than  to  the  dry  season  form.  Of  the  two  females,  one 
is  intermediate,  as  in  the  male,  and  the  other  of  the  normal  wet  season  form. 

DISTRIBUTION.  Throughout  Burma,  except  in  the  extreme  south  of  Tenasserim;  Thailand. 

Genus  LOGANIA  Distant 

Logania  Distant,  1884:  197,  208.  Type-species:  Logania  malayica  Distant,  1884:  208,  pi.  22,  fig.  21  $,  by 

monotypy.  Gender  feminine. 
Malais  Doherty,  1889: 414, 415, 436.  Type-species:  Loganiasriwa Distant  sensu  Doherty,  1889  [=  Logania 

marmorata  Moore,  1884],  by  designation  of  Corbet,  1940a:  111.  Gender  feminine.  [Synonymised  by  de 

Niceville,  1890:  32.] 

Eyes  smooth.  Antennae  half  the  length  of  the  forewing  costa,  with  about  36  segments  in  the  type-species; 


58  J.   N.  ELIOT 

shorter,  with  under  30  segments  in  L.  waltraudae;  rather  longer,  with  40-45  segments  in  the  marmorata- 
group  (=  Malais}.  Nudum  extending  widely  to  the  base  of  the  shaft.  Legs  much  shorter  than  in  Allotinus, 
with  the  tibiae  outwardly  swollen  and  the  fore-tarsi,  except  in  L.  waltraudae,  gradually  incrassate.  In  L. 
malayica  and  L.  waltraudae  the  male  fore-tarsus  ends  in  a  rather  long,  tapered,  down-curved  point,  but  in 
the  marmorata-group  the  fore-tarsus  ends  abruptly  in  a  rounded  pad  from  which  a  minute  point  is  directed 
downwards,  as  in  Allotinus.  Labial  palpi  shorter  than  in  Allotinus,  with  the  third  segment  usually  shorter 
than  one-half  of  the  second  segment  in  males,  but  may  be  slightly  longer  than  half  in  females.  Males  of  all 
species  have  a  small  double  hair  tuft  on  the  sternum  of  the  eighth  abdominal  segment.  The  type-species  has 
the  forewing  apex  produced  to  a  sharp  point  and  veins  MI  and  R5  have  a  long  common  stalk;  all  the 
remaining  species  have  a  rounded  or  square  apex  and,  except  in  L.  nehalemia,  the  stalk  of  veins  Mj  and  R5 
is  absent  or  short.  There  is  no  trace  of  a  humeral  vein  on  the  hindwing.  Males  of  the  ma/ay/ca-group  have 
vein  A/3  of  the  forewing  unswollen;  in  the  marmorata-group  the  basal  portion  of  vein  M3  is  briefly  swollen 
and  clothed  with  small,  specialised  scales  which  are  about  the  same  size  as  those  of  the  nivalis-group  of 
Paragerydus.  In  L.  regina,  and  probably  also  in  L.  paluana,  the  swelling  is  inconspicuous  and  partly  hidden 
by  normal  cover  scales.  The  underside  pattern  is  generally  similar  to  that  of  Allotinus,  but  the  ground 
colour  is  seldom  uniform,  being  mottled  in  shades  of  black,  brown  and  white,  for  which  reason  Evans 
coined  the  popular  name  'Mottles'  for  the  genus.  On  the  forewing  the  usual  lycaenid  markings  may  be 
difficult  to  make  out,  but  on  the  hindwing  they  are  usually  apparent  and  the  postdiscal  series  may  be 
catenulate.  In  some  species  there  is  very  great  individual  variation  in  the  extent  of  whitish  scaling  on  the 
upperside  of  the  hindwing,  which  may  be  absent  or  cover  almost  the  whole  of  the  wing.  The  male  genitalia 
are  of  the  usual  miletine  type  and  are  rather  constant  in  appearance,  except  that  L.  waltraudae  shows  some 
characters  suggestive  of  Allotinus. 

Doherty  (1889:  414,  415)  erected  the  generic  name  Malais  and  wrote  that  'it  will  include  L.  marmorata 
and  L.  sriwa  (probably  the  same  species)  and  one  or  two  rare  kinds  undescribed'.  He  had  before  him  only  a 
single  female  fronrMergui  which  he  treated  with  some  doubt  as  the  same  as  the  taxon  named  Logania  sriwa 
Distant,  1886.  De  Niceville  (1890:  33),  who  had  custody  of  the  type  of  L.  marmorata  Moore,  1884,  stated 
that  he  had  examined  Doherty's  female  of  'sriwa'  and  found  that  'it  appears  to  differ  from  L.  marmorata 
only  by  the  greater  prominence  of  all  the  markings  of  the  underside'.  As  the  females  of  L.  sriwa  and  L. 
marmorata  differ  so  strongly  it  is  inconceivable  that  Doherty's  female  was  really  sriwa,  and  highly  probable 
that  de  Niceville  was  correct  in  assuming  that  it  was  L.  marmorata.  I  can  find  no  convincing  evidence  that  L. 
sriwa  has  ever  been  taken  in  Burma,  although  it  was  recorded  by  Evans  (1932)  and  Cantlie  (1963)  from 
Mergui,  probably  on  the  basis  of  Doherty's  misidentified  female. 

Soon  afterwards  Doherty  (1891o:  29)  doubted  if  Malais  was  distinct  from  Logania;  and  though  he  did  not 
formally  synonymise  the  two  he  thereafter  used  Logania  for  species  which  would  fall  naturally  into  Malais. 
Bingham  (1907:  302),  Swinhoe  (1910:  200),  Evans  (1932:  199)  and  Cantlie  (1963:  2)  all  treated  Malais  as  a 
subjective  synonym  of  Logania,  as  did  Corbet  (1940a:  111)  who  stated  that  L.  sriwa  was  its  type-species. 
However,  as  Doherty's  'sriwa'  was  misidentified,  Corbet's  statement  cannot,  under  Article  70  of  the  Code, 
be  held  to  constitute  a  valid  type  selection.  Fruhstorfer  (1914;  1915),  whilst  using  Logania  as  the  generic 
name,  employed  Malais  in  a  subgeneric  sense  for  the  'Artengruppe'  with  banded  legs  and  rounded 
forewing  apex  which  includes  L.  marmorata  and  L.  sriwa.  Hemming  (1960:  11),  apparently  unaware  of 
Corbet's  action,  designated  Logania  malayica  Distant,  1884,  as  type-species  of  Malais.  His  action  is 
invalid,  since  malayica  was  not  one  of  the  species  originally  included  in  Malais  by  Doherty;  indeed,  the 
latter  (1889:  437)  had  specifically  excluded  malayica  from  his  new  genus,  stressing  that  it  was  a  true 
Logania.  Malais  remains,  therefore,  a  valid  and  available  genus-group  name,  which  can  be  used  as  a 
subgenus,  largely  in  the  sense  in  which  it  was  employed  by  Fruhstorfer,  by  those  authors  who  consider  that 
the  differences  between  Logania  and  Malais  are  greater  than  those  between  species-groups.  Malais  is, 
however,  still  without  a  properly  established  type-species,  and  any  author  intending  to  use  it  will  have  to 
refer  the  matter  to  the  International  Commission  on  Zoological  Nomenclature,  as  required  under  Article 
70  of  the  Code,  with  a  recommendation  that  Logania  marmorata  Moore,  1884,  be  designated  as 
type-species. 

The  genus  ranges  from  Peninsular  India  through  the  Archipelago  to  New  Guinea  and  the  Bismarcks,  and 
comprises  10  species. 

Key  to  the  species  of  Logania 

1  cf  upperside  of  forewing  with  vein  M3  unswollen  and  clothed  with  normal  cover  scales,  cf 

fore-tarsus,  so  far  as  known,  ending  in  a  tapered,  down-curved  point 2 

Cf  upperside  of  forewing  with  basal  part  of  vein  M3  swollen  and  clothed  with  specialised  scales. 

Cf  fore-tarsus  ending  abruptly ,  but  with  a  small  point  directed  downwards  from  its  lower  edge  4 

2  Forewing  apex  not  produced 3 


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59 


Forewing  apex  produced  to  a  sharp  point malayica  (p.  59) 

3  Upperside  of  hindwing  all  white waltraudae  (p.  61) 

Upperside  of  hindwing  with  outer  half  black  and  inner  half  white nehalemia  (p.  60) 

4  Legs  banded,  with  a  specially  prominent,  broad,  brown  band  on  outer  half  of  tibiae 5 

-  Legs  freckled,  not  clearly  banded 8 

5  $  upperside  with  grey  to  white  areas  except  in  marmomta  diehli 6 

-  $  upperside  entirely  brown obscura  (p.  68) 

6  Forewing  very  weakly  crenulate .  Underside  of  hindwing  with  a  white  streak  or  patch  bearing  few 

if  any  striae 7 

-  Forewing  termen  crenulate.  Underside  of  hindwing  without  an  unstriated  area...     marmorata  (p.  64) 

7  Upperside  of  hindwing  pale  grey  to  whitish regina  (p.  62) 

-  Upperside  of  hindwing  white  with  a  broad,  black  border paluana  (p.  63) 

8  Forewing  termen  only  very  weakly  crenulate 9 

-  Forewing  termen  crenulate watsoniana  (p.  73) 

9  Underside  mottling  more  or  less  ochreous distant!  (p.  69) 

-  Underside  mottling  dark  brown;  no  ochreous  tinge hampsoni  (p.  72) 

Logania  malayica  Distant 
(Figs  38,  39,  cf  genitalia) 
Logania  malayica  Distant,  1884:  208. 

The  species  is  instantly  recognisable  by  the  pointed  and  produced  forewing  apex.  The  sexes  are  alike, 
above  white  with  a  blackish  forewing  border  expanding  from  less  than  1-0  mm  at  the  tornus  to  nearly 
mid-costa.  Underside  white  densely  mottled  with  reddish  brown  striae.  The  legs  are  buff-brown  freckled 
with  whitish  scales,  and  the  male  fore-tarsus  ends  in  a  comparatively  long,  tapered,  down-curved  point. 

The  species  occurs  from  Peninsular  Thailand  to  Malaya,  Sumatra,  Borneo  and  the  Philippines.  There 
are  two  subspecies  with  somewhat  different  male  genitalia. 

Key  to  the  subspecies  of  L.  malayica 

1    Underside  irregularly  mottled  and  blotchy,  with, some  of  the  usual  lycaenid  markings  apparent 

malayica  malayica  (p.  60) 

-  Underside  regularly  and  densely  mott;      ,  with  none  of  the  lycaenid  markings  apparent 

malayica  subura  (p.  60) 


Fig.  38    Logania  malayica  malayica  Distant;  Malay  Peninsula.  Male  genitalia. 


60  J.  N.  ELIOT 

Logania  malayica  malayica  Distant 

(Fig.  38,  cf  genitalia) 

Logania  malayica  Distant,  1884:  208,  pi.  22,  fig.  21  $,  text-fig.  61  hind-leg.  Holotype  $,  WEST  MALAYSIA: 

Sungei  Ujong  (not  located). 
Logania  malayica  malayica  Distant;  Fruhstorfer,  1914:  23;  1916:  805,  pi.  141f  cf ;  Corbet,  1940a:  111,  fig.  1 

Cf  valva;  Fleming,  1975:  22,  pi.  58,  fig.  L51  $;  Eliot,  1978:  241,  pi.  20,  fig.  8  cf . 

On  the  underside  the  mottling  is  irregular  and  coalesced  into  blotches  in  places,  and  covers  both  wings 
except  for  a  white  area  above  the  forewing  dorsum  in  spaces  la  and  Ib.  The  usual  lycaenid  markings  can  be 
partly  made  out  with  difficulty. 

DISTRIBUTION.  Peninsular  Thailand;  West  Malaysia;  Sumatra;  Borneo,  including  Pulo  Laut. 

Logania  malayica  subura  Fruhstorfer 
(Fig.  39,  cf  genitalia) 

Logania  malayica  Distant;  Semper,  1889: 160,  pi.  31,  fig.  3  cf. 

Logania  malayica  subura  Fruhstorfer,  1914:  23;  1916:  805.  Holotype  cf ,  PHILIPPINES:  Mindanao  (SM). 

On  the  underside  the  mottling  of  striae  is  regular  and  without  blotches;  none  of  the  usual  lycaenid  markings 
can  be  made  out.  On  the  forewing  the  white  area  above  the  dorsum  extends  into  space  2.  The  male  genitalia 
differ  more  than  usual  in  subspecies,  and  it  may  be  that  subura  has  achieved  species  status. 

Fruhstorfer  named  subura  from  Semper's  figure,  so  the  male  depicted  therein  is  automatically  the 
holotype. 

DISTRIBUTION.  Probably  throughout  the  southern  Philippines,  but  I  have  only  seen  examples  from 
Mindanao  and  Samar. 


Fig.  39    Logania  malayica  subura  Fruhstorfer;  Samar.  Male  genitalia. 

Logania  nehalemia  Fruhstorfer  stat.  rev. 
(Figs  40,  cf  genitalia;  78  cf ) 

Logania  nehalemia  Fruhstorfer,  1914:  25;  1916:  808.  Holotype  '$'  recte  cf,  NEW  GUINEA  (BMNH) 

[examined]. 
Logania  hampsoni  nehalemia  Fruhstorfer;  D'Abrera,  1971:  384. 

Above,  the  forewing  is  white  with  a  black  border  which  curves  in  above  the  cell  to  the  wing  base;  the 
hindwing  has  the  basal  half  white  and  the  outer  half  black.  The  underside  is  white,  with  dense,  dark  brown 
striae  more  or  less  corresponding  with  the  areas  which  are  black  on  the  upperside. 
In  the  unique  male  holotype,  which  has  hitherto  been  regarded  as  a  female,  the  legs  are  missing  except 


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61 


Fig.  40    Logania  nehalemia  Fruhstorfer;  New  Guinea.  Male  genitalia. 

for  the  femur  and  tibia  of  one  hindleg  with  the  scales  rubbed  off;  these  hardly  differ  from  the  femur  and 
tibia  of  L.  malayica.  The  male  genitalia  are  chiefly  distinguished  by  the  phallus,  which  is  considerably 
stouter  than  that  of  its  congeners  except  for  L.  waltraudae. 

The  holotype  is  labelled  /Type  [red]/New  Guinea.  Hewitson  Coll.  79-69  Miletus  1/9  holotype  Logania 
nehalemia  Fruhst.  [in  Corbet's  hand]/.  Given  its  early  date  of  capture,  the  type-locality  is  likely  to  lie  in  the 
north-western  part  of  Irian  Jay  a. 

DISTRIBUTION.  New  Guinea. 

Logania  waltraudae  sp.  n. 

(Figs  41,  cT  genitalia;  107  cf ) 

C?  forewing  10-0  mm.  Generally  similar  in  appearance  to  the  sympatric  L.  malayica  subura,  and  like  it  with 
the  basal  portion  of  vein  M3  unswollen  and  clothed  with  normal  cover  scales;  but  differing  in  having  the 
forewing  apex  rounded  and  veins  MI  and  R5  connate. 


Fig.  41     Logania  waltraudae  sp.  n. ;  Samar.  Male  genitalia. 


62  J.  N.  ELIOT 

Upperside  white;  forewing  with  a  blackish  brown  apical  border  running  from  just  below  vein  Cu2  on  the 
termen  to  just  above  the  cell  apex  on  the  costa;  hindwing  with  a  dark  brown  marginal  hairline.  Cilia  dark 
brown.  Underside  pale  brown  very  densely  mottled  with  darker  brown  striae  except  on  the  forewing  in 
most  of  spaces  Ib  and  2  and  in  the  basal  part  of  space  3,  which  are  white.  A  dislocated  series  of  postdiscal 
spots  can  just  be  made  out  on  the  forewing. 

Antennae  just  under  half  the  length  of  the  forewing  costa,  thinner  than  in  L.  malayica,  with  probably 
28-30  segments  (both  antennal  clubs  are  broken  off  just  before  the  tip  after  26  segments).  The  middle  shaft 
segments  are  just  over  twice  as  long  as  wide  (in  the  remaining  Logania  species  these  segments  are  nearly  as 
wide  as  long).  The  nudum  extends  widely  to  the  base  of  the  shaft,  which  is  brown  on  the  upper  surface  with 
a  central  buff  patch  on  each  segment.  The  palpi  are  clothed  with  brown  and  a  few  buff  adpressed  scales  and 
are  exceptionally  short,  not  protruding  beyond  the  head,  with  the  third  segment  comparatively  stout  and 
only  a  quarter  the  length  of  the  second  segment.  The  legs  are  about  as  long  as  those  of  L.  malayica,  which 
they  resemble  in  having  the  fore-tarsus  ending  in  a  tapered,  down-curved  point;  but  they  differ  in  several 
respects.  The  femora,  tibiae  and  tarsi  are  subequal;  the  femora  are  broader  and  somewhat  flattened;  the 
foretibiae  are  narrow  and  cylindrical,  and  the  middle  tibiae  are  slightly  swollen,  the  swelling  being  greatest 
in  the  upper  half.  The  hind  legs  are  missing.  Body  dark  brown,  slightly  paler  beneath;  the  abdominal  hair 
tufts  on  the  eighth  sternum  are  smaller  than  in  the  other  species  of  Logania.  The  male  genitalia  are  broadly 
of  Logania  type,  but  the  phallus  is  distinctive,  while  the  triangular  vinculum  flap  and  strut  running  parallel 
to  the  lower  edge  of  the  uncus  plate  recall  those  structures  in  Allotinus. 

$  unknown. 

MATERIAL  EXAMINED 

Holotype  cf ,  Philippines:  Samar,  18.viii.l980(C.  G.  Treadaway)  (coll.  Treadaway,  but  will  be  deposited 
in  due  course  in  SM). 

The  species,  which  is  named  after  Mrs  Treadaway,  occupies  an  isolated  position  in  the  genus,  differing 
from  the  remainder  in  palpi,  antennae,  legs  and  male  genitalia,  and  possibly  deserves  to  be  placed  in  a 
separate  subgenus. 

Logania  regina  (H.  Druce) 

(Fig.  42,  cf  genitalia) 
Miletus  regina  H.  Druce,  1873:  348. 

This  and  subsequent  species  differ  from  the  foregoing  species  in  the  male  fore-tarsus,  which  is  not  tapered 
but  ends  abruptly  in  a  rounded  pad  from  the  lower  side  of  which  a  small,  short  point  is  directed  downwards, 
as  in  Allotinus.  The  legs,  in  this  and  the  next  three  species,  are  banded,  most  prominently  on  the  tibiae, 
with  whitish  and  brown,  while  in  the  remaining  species  they  are  freckled  and  sometimes  have  obscure 
longitudinal  streaks. 

A  distinguishing  character  of  L.  regina  is  the  white  ground  colour  of  the  underside  of  the  hindwing, 
which  is  almost  devoid  of  striae  in  a  streak-like  area  running  from  the  wing  base  to  the  termen  through  the 
upper  part  of  the  cell  and  space  6. 

On  the  upperside  the  male  has  a  fuscous  border  on  the  forewing  tending  to  run  narrowly  along  the  costa 
to  the  wing  base,  and  a  fuscous  costal  area  above  vein  6  on  the  hindwing.  The  rest  of  the  wings  are  whitish  to 
bluish  grey.  The  female  has  a  narrower  forewing  border  and  the  pale  areas  on  both  wings  are  whiter  than  in 
the  male. 

In  the  female  of  this  species  and  of  L.  paluana  (infra)  the  abdomen  is  longer  than  in  the  other  Logania 
species,  and  extends  just  beyond  the  hindwing  tornus. 

The  species  has  a  restricted  distribution  in  Sundaland,  excluding  Java,  Palawan  and  the  islands  off  the 
west  coast  of  Sumatra,  but  has  reached  the  Sulu  Is.  where  it  must  be  a  recent  immigrant.  There  are  two 
subspecies. 

Key  to  the  subspecies  of  L.  regina 

1  cf  upperside  of  forewing  with  black  border  comparatively  wide,  filling  whole  of  space  5.  $ 
forewing  border  reaching  dorsum;  underside  of  forewing  with  a  white  area  above  dorsum 
usually  reaching  vein  M2 regina  regina  (p.  63) 

-  cf  upperside  of  forewing  with  black  border  narrower,  not  reaching  base  of  space  5.  $  forewing 
border  fades  out  at,  or  just  before,  tornus;  underside  of  forewing  without  a  white  area 

regina  sriwa  (p.  63) 


THE  MILETINI  63 


Fig.  42    Logania  regina  regina  (H.  Druce);  Borneo.  Male  genitalia. 

Logania  regina  regina  (H.  Druce) 

(Fig.  42,  cf  genitalia) 

Miletus  regina  H.  Druce,  1873:  348,  pi.  32,  fig.  4  cT.  Holotype  cT,  BORNEO  (BMNH)  [examined]. 
Logania  regina  regina  (H.  Druce);  Fruhstorfer,  1914:  23;  1916:  806. 

Logania  evora  Fruhstorfer,  1916:  806.  Holotype  $,  PHILIPPINES:  'Sula  Is.'  recte  Sulu  Is  (SM)  [examined] 
Syn.  n. 

In  the  male  most,  if  not  all,  the  pale  areas  on  the  upperside  are  clothed  with  bluish  grey  scales.  The  female  is 
much  whiter,  with  a  narrower  forewing  border  which  is  usually  about  1-5  mm  wide  at  the  tornus.  On  the 
underside  of  the  forewing  there  is  sometimes  a  narrow  whitish  area  along  the  dorsum,  the  corresponding 
area  in  the  female  being  much  wider  and  usually  extending  into  the  basal  half  of  space  4. 

DISTRIBUTION.  Borneo;  Sulu  Is  (only  known  from  unique  holotype).  In  BMNH  there  is  a  single  male  from 
Pulo  Laut,  in  which  the  fore  wing  border  is  even  narrower  than  in  subsp.  sriwa,  while  the  pale  areas  are 
white,  so  that  the  general  appearance  is  of  a  female;  it  probably  represents  a  further  subspecies. 

Logania  regina  sriwa  Distant 

Logania  sriwa  Distant,  1886a:  531;  18866:  452,  pi.  44,  fig.  16  $;  Evans,  1932:  212.  Holotype  $,  WEST 

MALAYSIA  (not  located). 
Logania  regina  sriwa  Distant;  Fruhstorfer,  1914:  23;  1916:  805,  pi.  141f  $;  Corbet,  1940«:  112,  fig.  2  cf 

valva;  Cantlie,  1967:  28;  Fleming,  1975:  22,  pi.  58,  fig.  L52  ?;  Eliot,  1978:  241,  pi.  20,  fig.  9  cf . 

The  male  is  usually  whiter,  with  a  narrower  border,  than  nominate  regina.  In  the  female  the  forewing 
border  usually  fades  out  at  or  before  the  tornus.  On  the  underside  both  sexes  lack  a  white  area  above  the 
forewing  dorsum,  and  on  the  hindwing  the  white  streak  is  narrower. 

DISTRIBUTION.  West  Malaysia;  Sumatra;  peninsular  Thailand  (Pinratana,  1981). 

Logania  paluanasp.  n. 

(Fig- 79  9) 

This  taxon,  at  present  only  known  from  two  females,  appears  to  replace  L.  regina,  of  which  it  may  be  a 
subspecies,  in  Sulawesi.  It  is  larger,  with  forewing  length  17-0  mm,  compared  with  an  average  of  14-0  mm  in 
regina,  and  differs  additionally  as  follows.  On  the  upperside  the  forewing  border  is  narrower,  ending 
beyond  the  middle  of  the  costa.  On  the  hindwing  there  is  a  black  border  measuring  nearly  4-0  mm  at  the 
dorsum,  expanding  to  4-5  mm  at  vein  Cu2  and  thence  decreasing  to  1-0  mm  at  the  apex  whence  it  is 
continued  as  a  blackish  line  along  the  costa  to  the  wing  base.  The  underside  is  generally  marked  as  in 


64  J.  N.  ELIOT 

regina,  but  on  the  hindwing  the  comparatively  well-defined  white  streak  ofregina  is  replaced  by  a  wider  and 
more  obscure,  because  more  heavily  striated,  white  discal  patch  extending  to  vein  Cu\  and  the  lower  edge 
of  the  cell.  The  postdiscal  markings  are  olive-brown  and  there  is  a  well-defined  marginal  olive-brown  line 
0-5  mm  wide  on  the  forewing  and  between  veins  Cu2  and  M2  on  the  hindwing. 

MATERIAL  EXAMINED 

Holotype  9,  Sulawesi:  labelled  /G.  Rangkoean,  Paloe,  West  Celebes,  900',  Nov.  1936  (J.  P.  A.  Kalis) 
(BMNH). 

Paratype.  1  9 ,  data  as  holotype  (BMNH). 

Logania  marmorata  Moore 

(Fig.  43,  cf  genitalia) 
Logania  marmorata  Moore,  1884:  22. 

The  species  can  be  recognised  by  its  crenulate  forewing  termen  and  banded  legs,  which  are  shorter  and 
stouter  than  in  any  other  Logania  species.  Except  in  the  subspecies  from  Simeulue,  both  sexes  always  have 
a  pale  area  on  the  forewing,  which  is  more  extensive  in  the  female,  but  in  the  male  of  some  subspecies  it  is 
reduced  to  a  small  discal  patch.  The  hindwing  of  the  male  is  normally  brown,  at  least  in  the  wet  season, 
while  that  of  the  female  bears  some  grey  scales;  but  in  Nias  and  south  Sumatra  the  hindwing  is  partly  grey  in 
the  male  and  nearly  all  white  in  the  female. 

The  species  ranges  from  central  Burma  to  Vietnam  and  throughout  Sundaland,  the  Lesser  Sunda  Is.  and 
the  Philippines  into  north  Sulawesi.  The  dividing  line  between  subspecies  is  difficult  to  draw  because  of  the 
high  degree  of  individual  variation;  10  are  provisionally  recognised. 

Key  to  the  subspecies  of  L.  marmorata 

1  9  upperside  of  forewing  with  basal  half  whitish 2 

-  9  upperside  of  forewing  unmarked  brown marmorata  diehli  (p.  66) 

2  (if  upperside  of  hindwing  plain  brown 3 

-  cf  upperside  of  hindwing  partly  white  or  grey 11 

3  cf  upperside  of  forewing  with  some  bluish  grey  scaling  reaching  wing  base 4 

-  cf  upperside  of  forewing  with  brown  wing  base 6 

4  cf  upperside  of  forewing  with  pale  area  outwardly  white  but  wing  base  and  space  la  rather 

dark,  having  many  brown  scales  intermixed  with  the  grey.   9  upperside  of  hindwing 
sometimes  plain  brown 5 

-  cf  upperside  of  forewing  with  pale  area  outwardly  white  becoming  light  bluish  grey  at  wing 

base  and  in  space  la.  $  upperside  of  hindwing  with  at  least  some  grey  scaling  below  vein  6 

marmorata  damis  (p.  65) 

5  cf  upperside  of  forewing  with  white  extending  fully  across  space  Ib.  $  underside  of  forewing 

without  a  whitish  area  beyond  cell marmorata  hilaeira  (p.  65) 

-  cf  upperside  of  forewing  with  white  area  not  below  midspace  Ib.  $  underside  of  forewing  with 

a  whitish  area  beyond  cell  extending  from  vein  A\  to  veins  M2  or  MI 

dry  season  form  of  marmorata  javanica  (p.  67) 

6  9  underside  of  forewing  without  a  prominent  whitish  area  beyond  the  cell 7 

9  underside  of  forewing  with  a  broad  whitish  area  beyond  the  cell  stretching  from  vein  A\  to 

vein  M2. 

Cf  upperside  of  forewing  with  an  ovate  discal  white  patch  about  2-5  mm  wide  at  base  of 
spaces  4,3,2  and  sometimes  just  entering  space  Ib marmorata  munichya  (p.  66) 

7  cf  upperside  of  forewing  with  discal  white  patch  at  least  2-0  mm  wide,  reaching  and  often 

crossing  vein  Cu2.  $  upperside  of  forewing  with  base  more  or  less  grey-scaled 8 

-  cf  upperside  of  forewing  with  discal  patch  about  1-25  mm  wide  and  not  below  mid-space  2.  9 

upperside  of  forewing  with  base  brown,  bearing  only  a  trace  of  grey  scaling 

wet  season  form  of  marmorata  javanica  (p.  67) 

8  Cf  9  underside  comparatively  paler  and  browner ;  base  and  disc  of  forewing  not  blackish 9 

Cf  9  underside  dark;  base  and  disc  of  forewing  blackish 10 

9  cf  upperside  of  forewing  with  white  patch  usually  not  below  vein  Cu2.  Underside  comparative- 

ly dark.  Continental wet  season  form  of  marmorata  marmorata  (p.  65) 

Cf  upperside  of  forewing  with  white  patch  usually  crossing  vein  Cu2.  Underside  comparatively 
light.  Palawan marmorata  palawana  (p.  67) 


THE  MILETINI  65 

10  9  upperside  of  hindwing  with  some  grey  scaling marmorata  samosata  (p.  67) 

-  9  upperside  of  hindwing  brown marmorata  Faustina  (p.  68) 

11  cf  upperside  of  hindwing  brown  streaked  with  grey  below  vein  MI.  9  upperside  of  hindwing 

mostly  dirty  whitish  below  vein  MI dry  season  form  of  marmorata  marmorata  (p.  65) 

-  cf  upperside  of  hindwing  white  below  vein  M\  except  for  an  inwardly  diffuse  brown  border 

1 '0-1 -5  mm  wide.  9  upperside  of  hindwing  nearly  all  white marmorata  lahomius  (p.  66) 

Logania  marmorata  marmorata  Moore 

Logania  marmorata  Moore,  1884:  22;  1886:  39,  pi.  3,  fig.  7;  de  Niceville,  1890:  33,  frontispiece,  fig.  128  $ 
holotype;  Bingham,  1907:  303;  Swinhoe,  1910:  200,  pi.  618,  figs  1,  la  'cf'  recte  <j>;  Evans,  1932:  213. 
Holotype  $,  BURMA:  Mergui  (ZSI). 

[Malaissriwa  (Distant)  sensu  Doherty,  1889:  436.  Misidentification.] 

Logania  marmorata  marmorataMoore;  Fruhstorfer,  1914: 23;  1916:  806,  pi.  141f  cf  $ ;  Corbet,  19400: 112; 
Cantlie,  1963:  29. 

The  subspecies  occurs  in  two  seasonal  forms,  at  least  in  the  northern  part  of  its  range.  In  the  wet  season 
form  both  wings  are  brown,  with  a  small  white  patch  on  the  forewing  above  vein  Cu2  in  the  male  and  a  much 
larger  white  area  which  is  greyish  basally  in  the  female.  In  the  dry  season  the  white  areas  of  the  forewing  are 
much  enlarged  in  both  sexes,  the  hindwing  is  more  or  less  overlaid  below  vein  MI  with  grey  or  whitish 
scales,  especially  in  the  female,  and  on  the  underside  of  the  forewing  there  is  no  whitish  area  beyond  the 
cell.  Intermediate  season  examples  may  be  indistinguishable  from  subsp.  damis. 

DISTRIBUTION.  Burma,  from  Karen  Hills  to  Mergui;  Thailand;  Vietnam. 

Logania  marmorata  damis  Fruhstorfer 

Logania  massalia  damis  Fruhstorfer,  1914:  24;  1916:  807.  LECTOTYPE  cf ,  SINGAPORE  (BMNH),  here 

designated  [examined]. 
Logania  marmorata  damis  Fruhstorfer;  Corbet,  1940a:  111;  Fleming,  1975:  22,  pi.  58,  fig.  L53  9;  Eliot, 

1978:  241,  pi.  20,  fig.  10  cf. 

In  the  wet  and  only  seasonal  form  the  pale  areas  on  the  forewing  are  whiter  and  more  extensive  than  in  any 
other  subspecies  except  lahomius.  In  the  male  the  pale  area  is  outwardly  white,  inwardly  rather  pale  bluish 
grey,  reaches  the  dorsum  and  fills  the  cell;  the  hindwing  is  brown.  In  the  female  the  pale  area  is  whiter  and 
broader,  and  the  hindwing  always  bears  at  least  some  grey  scales  below  vein  M\. 

I  designate  as  lectotype  of  damis  a  male  in  BMNH  labelled  /Type  [red]/Singapora  II.  95/almost  certainly 
type  of  Logania  massalia  damis  Fruhst.  [in  Corbet's  hand]/. 

DISTRIBUTION.  Peninsular  Thailand;  West  Malaysia;  Singapore;  east  coastal  region  of  Sumatra. 

Logania  marmorata  hilaeira  Fruhstorfer 

Logania  obscura  Distant  &  Pryer,  1887:  266.  Syntypes,  BORNEO:  Sandakan  (not  located).  [Secondary 

homonym  of  Logania  obscura  (Rober,  1886).] 
Logania  marmorata  hilaeira  Fruhstorfer,  1914:  23;  1916:  806;  Corbet,  1940a:  112.  LECTOTYPE  cf, 

SUMATRA  (BMNH),  here  designated  [examined]. 
Logania  marmorata  stenosa  Fruhstorfer,  1914:  23  (nomen  nudum);  1916:  806;  Corbet,  1940a:  112. 

LECTOTYPE  9,  BORNEO  (BMNH),  here  designated  [examined].  Syn.  n. 
Logania  marmorata  obscura  Distant  &  Pryer;  Fruhstorfer,  1914:  23. 
Logania  massalia  nada  Fruhstorfer,  1914:  24;  1916:  807.  LECTOTYPE  cf,  SUMATRA  (BMNH),  here 

designated  [examined].  [Synonymised  by  Corbet,  1940a:  112.] 
Logania  marmorata  cineraria  Fruhstorfer,  1916:  806;  Corbet,  1940a:  112.  [Replacement  name  for  Logania 

obscura  Distant  &  Pryer,  1887.]  Syn.  n. 
Logania  massalia  sora  Fruhstorfer,  1916:  807.  LECTOTYPE  cf,  BORNEO  (BMNH),  here  designated 

[examined].  Syn.  n. 
Logania  marmorata  sora  Fruhstorfer;  Corbet,  1940a:  112. 

Above,  the  male  has  a  slightly  wider  forewing  border  than  subsp.  damis,  and  the  base  of  the  wing  is  darker 
with  brown  scales  intermixed  with  the  grey.  The  female  generally  has  a  darker  hindwing  which  is  often 
without  any  grey  scales. 

I  designate  as  lectotype  of  hilaeira  a  male  in  BMNH  labelled  /Type  [red]/CMB  IV.94/Fruhstorfer  Coll. 


66  J.  N.   ELIOT 

B.M.  1933-131/marmorata  hilaeira  Frhst.  [in  Fruhstorfer's  hand]/.  A  female  labelled  /Type  [red]/Type 
[Fruhstorfer  orange]/Sumatra  Montes  Battak  ex  coll.  Fruhstorfer/marmorata  Selesseh  15.VII.94/Fruh- 
storfer  Coll.  B.M.  1933-131/$  Allotype  of  Logania  marmorata  hilaeira  Fruh.  [in  Corbet's  hand]/  is  a 
paralectotype. 

In  his  earlier  work  (1914)  Fruhstorfer  did  not  list  or  describe  Logania  marmorata  stenosa,  either  through 
an  oversight  or  lapsus  calami,  but  he  mentioned  stenosa  twice  by  comparison  with  his  sub-spp.  javanica  and 
samosata.  In  1916  he  gave  a  brief  description  and  type-locality,  so  that  the  name  dates  from  1916.  I 
designate  as  lectotype  of  stenosa  a  female  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Sintang  Dr.  Martin  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/Marmorata  stenosa  Fr.  [in  Fruhstor- 
fer's hand]/. 

I  designate  as  lectotype  of  nada  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/CMB 
X.94/Fruhstorfer  Coll.  B.M.  1933-131/massalia  nada  Frhst.  [in  Fruhstorfer's  hand]/. 

I  designate  as  lectotype  of  sora  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Nord-Borneo  ex  coll.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/massalia  sora  Fr.  [in  Fruhstorfer's 
hand]/. 

DISTRIBUTION.  Northern  and  western  Sumatra  at  least  as  far  south  as  Padang;  Borneo,  including  Pulo  Laut; 
Cagayan  Sulu. 

Logania  marmorata  lahomius  (Kheil) 

Miletus  lahomius  Kheil,  1884:  27,  pi.  5,  figs  28,  29  cf .  Syntypes,  NIAS  (not  located). 
Logania  regina  lahomius  (Kheil)  Fruhstorfer,  1914:  23. 
Logania  lahomius  (Kheil);  Fruhstorfer,  1916:  806. 
Logania  marmorata  lahomius  (Kheil);  Corbet,  1940a:  113. 

Above,  both  sexes  are  greyish  white  with  the  forewing  border  narrower  than  in  the  preceding  subspecies. 
The  hindwing  of  the  male  has  a  diffuse  fuscous  margin  about  1-5  mm  wide  and  the  costal  area  is  fuscous 
above  vein  MI,  but  in  the  female  the  margin  is  vestigial  or  absent  and  there  is  only  a  little  fuscous  dusting 
below  the  costa. 

DISTRIBUTION.  Nias  I.  There  is  a  pair  in  BMNH  taken  by  Doherty  at  Liwa,  in  the  extreme  south-west  of 
Sumatra,  which  differ  only  that  in  the  female  the  fuscous  scaling  below  the  hindwing  costa  is  solid  as  far  as 
vein  MI;  they  are  provisionally  placed  under  lahomius. 

Logania  marmorata  die/i/i  subsp.  n. 

Cf  upperside  brown;  forewing  with  a  small,  sullied,  circular  whitish  patch  2-0  mm  wide  surrounding  swollen 
portion  of  vein  M3  in  spaces  2,  3  and  4.  Underside  generally  pale,  with  postdiscal  markings  ill-defined; 
forewing  without  a  trace  of  a  white  or  paler  discal  area,  as  in  the  otherwise  rather  similar  subspecies  from 
Java. 

$  upperside  entirely  brown.  Underside  like  male. 

MATERIAL  EXAMINED 

Holotype  cf ,  Simeulue  (Simalur):  16-17. ii. 1984  (E.  Dieht)  (BMNH). 
Paratype.  1  $  (allotype),  data  as  holotype  (BMNH). 

The  subspecies  is  named  in  honour  of  the  captor,  Dr  Edvard  Diehl. 

It  is  instructive  that  the  two  extremes  of  geographical  variation  in  this  species  are  found  in  the 
neighbouring  islands  of  Nias  and  Simeulue,  the  former  having  the  most  extensive  white  markings,  the  latter 
the  least;  a  good  example  of  the  haphazard  course  of  evolution  in  small,  isolated  populations. 

Logania  marmorata  munichya  Fruhstorfer 

Logania  massalia  munichya  Fruhstorfer,  1914:  24;  1916:  807.  LECTOTYPE  cf ,  JAVA  (BMNH),  here 

designated  [examined]. 
Logania  marmorata  javanica  Fruhstorfer;  Corbet,  1940a:  112,  partim. 

In  this  and  the  remaining  subspecies  the  males,  at  least  in  the  dry  season,  have  a  brown  forewing  bearing  a 
white  discal  patch  and  without  grey  scales  at  the  wing  base.  In  munichya  the  white  patch  is  2-5  mm  wide  and 
lies  at  the  bases  of  spaces  4,  3  and  2  and  just  enters  space  Ib.  The  female  resembles  subsp.  hilaeira  on  the 
upperside  but  is  distinctive  in  possessing,  on  the  underside  of  the  forewing,  a  white  area  3-0-4-0  mm  wide 
beyond  the  cell  stretching  from  vein  A±  to  vein  M2. 


THE  MILETINI  67 

It  seems  likely  that  this  subspecies  and  the  east  Javanese  subsp.  javanica  may  represent  the  ends  of  a 
cline,  in  which  case  Corbet's  action  in  synonymising  munichya  with  javanica  would  be  justified. 

I  designate  as  lectotype  of  munichya  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Java  Occident.  Sukabumi  2000'  ex  coll.  Fruhstorfer/massalia  munichya  Frhst.  [in  Fruhstorfer's  hand]/. 

DISTRIBUTION.  West  Java. 

Logania  marmorata  javanica  Fruhstorfer 

Logania  marmorata  javanica  Fruhstorfer,  1914:  23;  1916:  806;  Corbet,  1940a:  112.  LECTOTYPE  cf ,  JAVA 

(BMNH),  here  designated  [examined]. 
Logania  massalia  glypha  Fruhstorfer,  1914:  23;  1916:  807.  LECTOTYPE  cf,  JAVA  (BMNH),  here 

designated  [examined].  [Synonymised  by  Corbet,  1940«:  112.] 
Logania  marmorata  Moore;  Piepers  &  Snellen,  1918:  18,  pi.  20,  fig.  21  cf  • 
[Logania  massalia  Doherty;  Piepers  &  Snellen,  1918:  18,  pi.  20,  figs  22a  cf ,  22b  $.  Misidentification.] 

The  male  differs  from  subsp.  munichya  in  the  smaller  white  patch  on  the  forewing,  which  is  1-25  mm  wide 
and  does  not  descend  below  mid-space  2.  The  female  differs  in  having  the  base  of  the  forewing  darker  and, 
on  the  underside,  in  lacking  the  prominent  white  patch  beyond  the  cell,  this  area  being  only  slightly  paler 
than  the  rest  of  the  wing. 

I  designate  as  lectotype  of  javanica  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 
Ostjava  Lawang  1897  ex  coll.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/Logania  javanica  Frhst.  [in 
Fruhstorfer's  hand]/.  Its  left  forewing  is  much  discoloured,  the  basal  two-thirds  of  the  wing  having  an  oily 
bluish  sheen;  the  right  forewing  is  undamaged.  The  figure  in  Piepers  &  Snellen  (1918),  showing  only  the  left 
half  of  a  male  with  a  basally  bluish  forewing,  was  obviously  made  from  the  lectotype;  Piepers  said  that  he 
had  himself  seen  no  Javanese  examples  of  L.  marmorata.  There  are  also  in  BMNH  two  females  of  the 
original  type-series  labelled  /Logania  javanica  Frhst.  [in  Fruhstorfer's  hand]/  which  are  paralectotypes. 

I  designate  as  lectotype  of  glypha  a  male  in  BMNH  labelled  /Type  [red]/Ostjava  Lawang  1897  ex  coll. 
Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/massalia  glypha  Frhst.  [in  Fruhstorfer's  hand]/. 

All  the  above  lectotypes  and  paralectotypes  appear  to  be  wet  season  forms.  There  is  in  BMNH  a  series  of 
4  cf ,  7  $  from  Sumbawa  labelled  /Sumbawa  Doherty  Sept  '91/  which  must  represent  the  dry  season  form 
and  which  I  provisionally  place  under  subsp.  javanica.  The  males  have  a  white  patch  on  the  forewing  similar 
to  that  of  subsp.  munichya,  but  the  base  of  the  wing  is  covered  with  rather  dark  bluish  grey  scales.  The 
females  also  are  rather  similar  on  the  upperside  to  that  sex  of  subsp.  munichya,  but  the  hindwing  is  paler 
brown  and  may  bear  some  grey  scales,  while  on  the  underside  of  the  forewing  the  whitish  area  beyond  the 
cell  is  narrower  and  more  sullied.  There  is  also  in  BMNH  a  single  female  labelled  /S.  Flores  xi.96.  Dry  s. 
Everett/  which,  despite  the  label  appears  to  be  a  wet  season  form  as  might  be  anticipated  from  the  date  of 
capture  at  the  change  of  seasons.  Above,  it  differs  only  slightly  from  females  of  subsp.  javanica  in  having 
the  base  of  the  forewing  browner,  with  only  a  very  few  overlying  grey  scales,  while  on  the  underside  of  the 
forewing  the  whitish  patch  beyond  the  cell  is  vestigial.  Provisionally  I  attach  it  also  to  subsp.  javanica. 

DISTRIBUTION.  East  Java;  Lesser  Sunda  Is.  (Sumbawa,  Flores). 

Logania  marmorata  palawana  Fruhstorfer 

Allotinus  (Logania)  distanti  Staudinger,  1889:  93,  pi.  1,  fig.  3  $.  Synt'ypes,  PALAWAN  (?  MNHU). 

[Secondary  homonym  of  Logania  distanti  Semper,  1889.] 
Logania  marmorata  palawana  Fruhstorfer,  1914:  23;  1916:  806.  [Replacement  name  for  Logania  distanti 

(Staudinger,  1889).] 

The  male  resembles  subsp.  munichya,  but  the  forewing  patch  is  a  little  larger  and  faintly  bluish  grey.  The 
female  is  most  like  subspp.  marmorata  and  hilaeira,  especially  on  the  underside,  but  on  the  upperside  of  the 
forewing  the  border  is  a  little  wider  at  the  tornus. 

Single  females  from  Luzon  in  BMNH  and  Marinduque  (coll.  Treadaway)  appear  to  belong  to  this 
subspecies. 

DISTRIBUTION.  Palawan;  Balabac  I.  (coll.  Treadaway);  Luzon;  Marinduque. 

Logania  marmorata  samosata  Fruhstorfer 

[Logania  obscura  Distant  &  Pryer  sensu  Semper,  1889:  160  partim,  pi.  31,  fig.  4  $.  Misidentification.] 
Logania  marmorata  samosata  Fruhstorfer,  1914:  23;  1916:  806.  Holotype  cf ,  PHILIPPINES:  Cebu  (SM), 
[examined]. 


68  J.  N.   ELIOT 

I  have  seen  only  two  females  and  no  males  from  Cebu.  The  former,  which  have  the  lower  part  of  the 
hindwing  lightly  grey-scaled,  differ  additionally  from  subsp.  palawana  by  a  much  darker  underside  with  the 
forewing  mostly  blackish.  A  female  from  Mindoro  is  similar. 

Fruhstorfer  named  the  subspecies  from  Semper's  fig.  4,  so  the  specimen  depicted  therein  is  automatically 
the  holotype. 

DISTRIBUTION.  Cebu;  probably  Mindoro. 

Logania  marmorata  Faustina  Fruhstorfer 
(Fig.  43,  cf  genitalia) 

[Logania  obscura  Distant  &  Pryer  sensu  Semper,  1889:  160  partim,  pi.  31,  fig.  5  $.  Misidentification.] 
Logania  marmorata  faustina  Fruhstorfer,  1914:  23;  1916:  806.  Holotype  9,  PHILIPPINES:  Mindanao  (SM), 
[examined]. 

All  the  females  I  have  seen  from  Mindanao,  Samar  and  Leyte  have  the  upperside  of  the  hindwing  plain 
blackish  brown;  otherwise  they  do  not  differ  from  subsp.  samosata.  The  males  resemble  subsp.  palawana 
on  the  upperside,  but  are  readily  separable  by  their  much  darker  blackish  undersides. 

Fruhstorfer  described  the  subspecies  from  Semper's  fig.  5,  so  the  female  from  Mindanao  depicted 
therein  is  automatically  the  holotype. 

DISTRIBUTION.  Mindanao;  Leyte;  Samar  (coll.  Treadaway);  Sulu  Is.:  Tawi  Tawi  (Tite,  1969). 


Fig.  43     Logania  marmorata  faustina  Fruhstorfer;  Mindanao.  Male  genitalia. 


Logania  obscura  (Rober) 
(Fig.  44,  cf  genitalia;  80  cT) 

Allotinus  obscurus  Rober,  1886:  52,  pi.  4,  fig.  8  cf .  Syntypes,  SULAWESI  (?  SMT). 

Allotinus  martinus  Fruhstorfer,  1913:  371;  1916:  814,  pi.  141h  $.  Holotype  $,  SULAWESI:  Buton  I. 
(BMNH)  [examined].  Syn.  n. 

Logania  donussa  Fruhstorfer,  1914:  24.  LECTOTYPE  $,  SULAWESI  (BMNH),  here  designated  [ex- 
amined]. Syn.  n. 

Logania  distanti  donussa  Fruhstorfer;  Fruhstorfer,  1916:  807. 

Logania  obscura  (Rober)  Fruhstorfer,  1916:  807. 

Logania  marmorata  obscurus  (Rober);  Corbet,  1940a:  112. 

In  the  past  L.  obscura  has  been  thought  to  replace  L.  marmorata  in  Sulawesi  and  its  satellite  islands,  and  it 
was  treated  as  a  subspecies  thereof  by  Corbet.  However,  the  very  recent  discovery  of  L.  marmorata  in 
north  Sulawesi  in  a  still  undescribed  subspecies  indicates  that  L.  obscura  is  a  distinct  species.  Its  status  as 
such  is  further  confirmed  by  the  following  characters.  The  female  is  all  brown,  whereas  the  male  has  a  white 
forewing  patch  -  a  reversal  of  the  usual  sexual  differences  in  Logania  marmorata  wherein  the  female  has 


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69 


Fig.  44     Logania  obscura  Rober;  Sulawesi.  Male  genitalia. 


more  extensive  white  areas  than  the  male  (except  in  Simeulue).  The  forewing  termen  and  apex  are  more 
rounded  and  the  crenulations  are  weaker.  The  legs  are  longer  and  thinner.  The  wing  span  is  considerably 
larger.  Finally,  there  is  a  small  difference  in  the  male  valva. 

The  holotype  of  martinus  has  no  head,  legs  nor  right  forewing.  It  bears  a  label  /damaged  in  shelter  by 
burst  pipe,  G.E.T./.  The  remaining  wings  show  no  significant  difference  to  normal  L.  obscura. 

I  designate  as  lectotype  ofdonussa  a  female  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/S. 
Celebes  Bua-Kraeng  5000'  Febr.  1896  H.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/Logania  donussa 
Fr.  [in  Fruhstorfer's  hand]/. 

A  single  male  from  Banggai  I.  in  coll.  Bedford  Russell  (Fig.  80)  has  the  forewing  patch  sullied  all  over 
with  fuscous  scales,  so  that  it  hardly  shows  up,  and  the  underside  is  darker  and  more  densely  striated  than  in 
typical  examples.  A  male  from  the  Toekan  Besi  Is.  in  BMNH  resembles  the  Banggai  male  on  the 
upperside,  but  on  the  underside  is  paler  than  typical  examples.  These  two  males  probably  represent  further 
minor  subspecies. 

DISTRIBUTION.  Sulawesi. 


Logania  distant!  Semper 
(Figs  45,  46,  cf  genitalia;  81  cf ) 
Logania  distanti  Semper,  1889:  161. 

In  this  species  I  combine  the  subspecies  which  have  hitherto  been  treated  as  pertaining  to  two  distinct 
species,  L.  distanti  and  L.  massalia.  Fruhstorfer,  and  also  Corbet  in  part,  confused  the  massalia 
subspecies-group  with  L.  marmorata,  but  the  two  species  are  easily  distinguished  by  several  characters. 
Firstly,  in  L.  distanti  the  legs  are  longer,  thinner  and  not  banded,  but  speckled  and  streaked  with  brown  on 
a  whitish  ground.  Secondly,  the  forewing  termen  is  barely  crenulate.  Thirdly,  the  underside  is  more 
ochreous  and  the  striations  are  usually  denser.  In  the  male  genitalia  the  phallus  is  much  less  strongly  bent. 

The  male  is  brown  above,  normally  with  a  whitish  or  bluish  grey  discal  patch  on  the  forewing,  but  in  the 
Philippines  and  Malay  Peninsula  (and  probably  elsewhere)  the  upperside  is  sometimes  unmarked.  On  the 
forewing  vein  M3  is  briefly  swollen  and  clothed  with  specialised  scales.  Females  show  very  great  individual 
variation,  the  forewing  white  or  very  pale  greyish  blue  with  a  broad  blackish  border,  while  the  hindwing, 
except  in  subsp.  staudingeri,  varies  from  brown  to  mainly  whitish  in  any  one  area. 

The  species  flies  from  India  to  the  Malay  Peninsula,  Sumatra,  Borneo  and  the  Philippines.  It  is 
apparently  absent  from  the  islands  off  the  west  coast  of  Sumatra  and  from  Java,  records  of  L.  massalia  from 
the  latter  island  applying  to  misidentified  L.  marmorata. 

Key  to  the  subspecies  of  L.  distanti 

1  cf  with  upperside  of  forewing  brown  to  brownish  at  base.  $  with  upperside  of  hindwing 
variable,  but  in  the  most  lightly  marked  examples  always  with  a  fuscous  border  1-0-2-0  mm 
wide  ..  2 


70  J.  N.   ELIOT 

Cf  upperside  of  fo rawing  with  pale  bluish  grey  area  reaching  wing  base .  $  upperside  of  hindwing 
all  white  except  for  sparse  fuscous  dusting  above  vein  A/i distant!  staudingeri  (p.  72) 

2  Cf  upperside  of  forewing  with  discal  patch,  if  present,  pale  bluish  grey 3 

Cf  upperside  of  forewing  with  discal  patch ,  if  present ,  white distant!  massalia  (p .  70) 

3  Philippine distant!  distanti  (p.  70) 

Bornean distanti  drucei  (p.  71) 

Logania  distanti  distanti  Semper 

(Fig.  81  Cf ) 

Logania  distanti  Semper,  1889:  161,  pi.  31,  figs  6,  7  $,  21  cf.  Holotype  <j>,  PHILIPPINES:  Cebu  (SM) 

[examined]. 

Logania  distanti  distanti  Semper;  Fruhstorfer,  1914:  24;  1916:  807. 
Logania  distanti  apsines  Fruhstorfer,  1914: 24;  1916:  807.  LECTOTYPE  cf ,  PHILIPPINES:  Mindanao  (SM), 

here  designated  [examined]. 
Logania  turdeta  Fruhstorfer,  1916:  807.  Holotype  'cf'  recte  $,  PHILIPPINES:  Cebu  (SM)  [examined]. 

Syn.  n. 

Semper  originally  described  L.  distanti  from  three  examples:  a  male  and  female  of  large  size  from 
Mindanao,  which  he  figured  at  figs  21  and  7  respectively,  and  a  small  female  from  Cebu  figured  at  fig.  6. 
Although  he  labelled  the  male  as  type  he  unfortunately  did  not  specify  it  as  such,  and  this  left  the  way  open 
for  Fruhstorfer  later  to  restrict  nominate  distanti  to  Cebu  and  to  name  the  Mindanao  pair  as  subsp.  apsines. 
Semper's  female  from  Cebu  is,  therefore,  automatically  the  holotype  of  distanti. 

Fruhstorfer 's  action,  taken  at  a  time  when  it  is  evident  that  he  had  not  even  seen  Semper's  specimens,  is 
doubly  regrettable,  since  a  year  later,  after  examining  Semper's  collection,  he  named  the  Cebu  female, 
which  he  then  inexplicably  mistook  for  a  male,  as  a  new  species,  L.  turdeta,  and  remarked  that  Semper  had 
confused  it  with  L.  distanti.  At  the  same  time  he  still  recorded  nominate  distanti  from  Cebu  and  maintained 
apsines  for  Mindanao  examples.  Semper's  original  female  from  Cebu,  which  is  the  holotype  of  both  distanti 
and  turdeta,  is  labelled  /CEBU/Coll.  C.  Semper/206/1004/No  6/Typus  [and  on  reverse]  Logania  turdeta 
(Fruh.)/.  The  small  ticket  reading  'No  6'  obviously  refers  to  Semper's  fig.  6.  The  specimen  has  extensive 
whitish  dusting  on  the  upperside  of  the  hindwing  and  agrees  fairly  well  with  the  butterfly  figured  by  de 
Niceville  (1894:  pi.  2,  fig.  13)  as  Logania  luca. 

Semper's  pair  from  Mindanao  may  or  may  not  be  a  valid  subspecies;  but  as  size,  by  which  alone 
Fruhstorfer  separated  them  as  subsp.  apsines,  is  an  unreliable  character.  I  treat  apsines  provisionally  as  a 
synonym  of  distanti.  Fruhstorfer  did  not  designate  a  type,  so  I  now  designate  the  male  as  lectotype;  it  is 
labelled  /Log.  Distanti  typ.  Semper/206/1004/16/Typus  [red]/.  The  small  ticket  reading  '16'  should  refer  to 
the  figure  in  Semper's  pi.  31;  but  in  fact  the  specimen  is  shown  at  fig.  21,  while  fig.  16  represents 
Parageryduspunctatus.  Possibly  the  labels  got  transferred  by  mistake,  as  in  other  cases  already  mentioned 
(p.  37)  during  Fruhstorfer's  examination  of  the  collection.  Above,  the  specimen  is  unmarked  brown,  but 
as  it  is  the  only  Mindanao  male  I  have  seen  it  would  be  premature  to  assume  that  an  all-brown  forewing  is  an 
unvarying  character  of  the  male  in  that  island. 

I  have  seen  no  males  from  Cebu,  but  have  examined  single  males  from  Luzon  and  Negros  kindly  sent  me 
by  Dr  Alistair  Ballantine  and  Mr  Yusuke  Takanami  respectively,  both  of  which  have  been  deposited  in 
BMNH.  The  former  is  as  large  as  the  lectotype  of  apsines,  but  has  a  small  diffuse  bluish  grey  discal  patch  on 
the  forewing.  The  latter  is  smaller,  and  has  a  more  extensive  bluish  grey  area  which  nearly  reaches  the  base 
of  the  forewing.  Judging  by  the  three  males  I  have  seen  I  expect  that  it  will  be  found  that  throughout  the 
Philippines  males  most  often  have  a  pale  discal  patch  on  the  forewing,  but  that  unmarked  males  may  occur 
anywhere  from  time  to  time,  just  as  they  do  in  the  Malay  Peninsula. 

Apart  from  the  two  females  in  Semper's  type-series  the  only  females  I  have  seen  are  two  from  Samar  in 
coll.  Treadaway.  Both  have  some  grey  dusting  on  the  upperside  of  the  hindwing,  but  in  neither  example  is  it 
as  extensive  as  in  the  holotype  of  distanti/ turdeta. 

DISTRIBUTION.  Probably  throughout  the  Philippines,  where  it  appears  to  be  everywhere  rare.  Examples 
seen  from  Luzon,  Negros,  Cebu,  Samar  and  Mindanao. 

Logania  distanti  massalia  Doherty  stat.  n. 

(Fig.  45,  cf  genitalia) 

Logania  massalia  Doherty,  18916:  37  '$'  recte  cT;  Bingham,  1907:  304;  Swinhoe,  1910:  202,  pi.  618.  figs  3, 
3b  cf ,  3a,  3c  '$'  recte  cf ;  Evans,  1932:  213.  Holotype  cf ,  INDIA:  Assam  (BMNH)  [examined]. 


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Fig.  45     Logania  dlstantl  massalia  Doherty;  Malay  Peninsula.  Male  genitalia. 


Logania  luca  de  Niceville,  1894:  28,  pi.  2,  fig.  13  <j>:  Evans,  1932:  213.  4  $  syntypes,  SUMATRA  and  WEST 

MALAYSIA  (probably  ZSI).  Syn.  n. 

Logania  luca  luca  de  Niceville;  Fruhstorfer,  1914:  24;  1916:  806. 

Logania  massalia  massalia  Doherty;  Fruhstorfer,  1914:  24;  1916:  807,  pi.  141f  cf ;  Corbet,  1940a:  112. 
Logania  massalia  luca  de  Niceville;  Corbet,  1940a:  112;  Fleming,  1975:  22,  pi.  58,  fig.  L54  cf  $;  Eliot, 

1978:  241,  pi.  20,  fig.  10  cf. 

Usually  the  male  has  a  circular  white  discal  patch  more  or  less  dusted  with  grey  scales  on  the  forewing,  but 
very  occasionally  the  patch  extends  to  the  wing  base  where  it  is  heavily  dusted  with  bluish  grey  and  fuscous 
scales,  as  in  single  males  in  BMNH  from  south  India  and  Burma  and  in  a  male  from  the  Malay  Peninsula  in 
coll.  Fleming.  Sometimes  the  white  patch  is  wanting  or  vestigial,  most  often  in  examples  from  the  Malay 
Peninsula.  The  hindwing  is  blackish  brown. 

Females  show  much  individual  variation,  and  cannot  be  reliably  separated  from  that  sex  of  the  nominate 
subspecies.  The  hindwing  may  be  unmarked  blackish  brown,  but  nearly  always  there  is  at  least  some  grey 
scaling.  De  Niceville  described  and  figured  as  L.  luca  females  from  Perak  and  Sumatra  in  which  the 
hindwing  was  more  than  half  whitish,  and  even  more  extreme  examples  with  paler  undersides  occur  in 
Sumatra  and  in  Assam  and  Burma  during  the  dry  season. 

In  BMNH  there  is  a  female  labelled  /Type  [red]/Perak  Jan. -Feb.  '90  W.  Doherty/Elwes  Coll.  1915-207/ 
L.  malais  Doh.  so  named  in  Elwes  Coll. /Perak  2c  Malais  sp.  undescribed  £ /Probably  not  a  type  but  agrees 
exactly  with  fig.  of  luca  Nic.  in  orig.  desc.  [in  Corbet's  hand]/.  In  fact  this  female,  which  may  conceivably  be 
one  of  de  Niceville's  four  female  syntypes,  does  not  agree  closely  with  de  Niceville's  figure,  since  the 
hindwing  has  hardly  any  whitish  scaling. 

DISTRIBUTION.  South  India  (cf ,  Thantipandal,  60  m.  NNW.  of  Madras);  Nepal;  north-east  India  (Sikkim  to 
Manipur);  Burma;  peninsular  Thailand;  West  Malaysia;  Sumatra. 

Logania  distant!  drucei  Moulton 

Logania  drucei  Moulton,  1911:  85,  fig.  9  cT;  Fruhstorfer,  1914:  24;  1916:  807.  Holotype  cf ,  BORNEO: 

Sarawak  (BMNH)  [examined]. 
Logania  massalia  drucei  Moulton;  Corbet,  1940a:  112. 

Probably  inseparable  from  subsp.  distanti,  but  as  I  have  seen  few  specimens  of  either  subspecies  I  maintain 
it  provisionally  as  distinct. 

The  only  males  I  have  seen  have  a  circular  bluish  grey  discal  patch  on  the  forewing  and  the  females  have  a 
brown  or  very  nearly  plain  brown  hindwing.  I  have  seen  no  females  similar  to  the  figure  of  luca. 

DISTRIBUTION.  Borneo,  excluding  Mt  Kina  Balu  where  it  is  replaced  by  subsp.  staudingeri. 


72  J.  N.   ELIOT 

Logania  distant!  staudingeri  H.  H.  Druce 
(Fig.  46,  cf  genitalia) 

Logan iastaudingeriH.  H.  Druce,  1895: 565,  pi.  31,  figs  13  cf ,  14  $ .  Syntypes,  BORNEO:  Mt  Kina Balu  (coll. 

Staudinger,  probably  in  MNHU). 

Logania  luca  staudingeri  Druce;  Fruhstorfer,  1914:  24;  1916:  806. 
Logania  marmorata  staudingeri  H.  H.  Druce;  Corbet,  1940a:  112. 

The  male  differs  from  subsp.  drucei  in  that  the  bluish  grey  patch  extends  to  the  base  and  dorsum  of  the 
forewing.  The  female  is  very  pale  bluish  grey,  with  the  usual  blackish  apical  and  marginal  border,  while  the 
hindwing  may  be  unmarked  except  for  some  fuscous  scaling  above  vein  Aft  and  a  marginal  blackish 
hairline,  or  there  may  be  a  diffuse  border  up  to  1-0  mm  wide. 

DISTRIBUTION.  Only  known  from  Mt  Kina  Balu. 


Fig.  46     Logania  distanti  staudingeri  H.  H.  Druce;  Borneo:  Kina  Balu.  Male  genitalia. 

Logania  hampsoni  Fruhstorfer 

(Fig.  47  cf  genitalia) 

Logania  hampsoni  Fruhstorfer,  1914:  25  'cf '  recte  $ ;  1916: 807.  LECTOTYPE  $ ,  NEW  GUINEA  (BMNH), 

here  designated  [examined]. 
Malais  meeki  Rothschild,  1915:  387.  LECTOTYPE  cf,  NEW  GUINEA:  Dampier  I.  (BMNH),  here 

designated  [examined].  Syn.  n. 
Logania  masana  Fruhstorfer,  1916:  808.  Holotype  cf,  NEW  GUINEA  (coll.  Staudinger,  probably  in 

MNHU).  Syn.  n. 

Logania  hampsoni  masana  Fruhstorfer;  D'Abrera,  1971:  384,  figs  cf  $. 
Logania  hampsoni  hampsoni  Fruhstorfer;  D'Abrera,  1971:  384. 

The  species  replaces  L.  distanti  to  the  east  of  Weber's  Line  and  is  rather  doubtfully  distinct.  It  differs  chiefly 
in  the  browner  underside  markings  and  striae,  which  lack  the  characteristic  ochreous  tone  of  distanti. 
Individual  variation,  so  conspicuous  a  feature  of  L.  distanti,  is  virtually  non-existent.  The  male  is  plain 
brown  above,  and  the  female  has  a  white  forewing  with  a  broad  brown  marginal  and  apical  border  and  a 
brown  hindwing. 

Fruhstorfer  described  L.  hampsoni  from  the  female,  which  he  mistook  for  the  male,  and  gave  as 
type-locality  Kumusi  River  in  Papua  New  Guinea.  As  there  were  no  specimens  in  his  collection  he  must 
have  described  the  species  from  memory  of  a  series  of  females  from  that  locality  which  he  had  seen  in  coll. 
Rothschild.  I  therefore  designate  as  lectotype  a  female  in  BMNH  labelled  /Lectotype  [purple]/Kumusi  R. 
N.  E.  Brit.  N.  Guin.  low  elev.  VIII-IX.07  (A.  S.  Meek)/Rothschild  Bequest  B.M.  1939-1/Lectotype  L. 
hampsoni  Fruhstorfer  'cf '  recte  $ ,  designated  by  J.  N.  Eliot  Nov.  1983/. 

Fruhstorfer's  error  over  the  sex  of  L.  hampsoni  was,  no  doubt,  responsible  for  the  mis-statement  by 


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Fig.  47     Logania  hampsoni  Fruhstorfer;  Papua  New  Guinea.  Male  genitalia. 

D'Abrera  (1971)  that  the  male  of  L.  hampsoni  hampsoni  has  a  white  basal  area  on  the  forewing,  but  can 
hardly  explain  his  further  mis-statement  that  the  type-locality  was  north-western  West  Irian. 

I  designate  as  lectotype  of  M.  meeki  a  male  in  BMNH  labelled  /Type  [red]/Dampier  Isl.  Feb.  &  March 
1914  (Meek's  Expedition)/Malais  meeki  Type  Rothsch./. 

DISTRIBUTION.  New  Guinea,  including  Dampier  I.;  New  Britain;  North  Moluccas:  Obi  I. 

Logania  watsoniana  de  Niceville  sp.  rev. 
(Fig.  48  cf  genitalia) 

Logania  watsoniana  de  Niceville,  1898:  143,  pi.  Z,  figs  17  cf ,  18  $;  Bingham,  1907:  303;  Swinhoe,  1910: 
201,  pi.  618,  figs  2,  2b  cf ,  2a,  2c  $  [intermediate  form];  Fruhstorfer,  1914:  23.  Syntypes,  BURMA:  North 
Shan  States  (probably  ZSI).  [Dry  season  form.] 

Logania  subfasciata  Tytler,  1915:  120.  Holotype  cf,  INDIA:  Manipur  (BMNH)  [examined].  [Wet  season 
form.]  [Synonymised  by  Cantlie,  1963:  29.] 

Logania  marmorata  watsoniana  de  Niceville;  Fruhstorfer,  1916: 806;  Corbet,  1940a:  112;  Cantlie,  1963: 29. 

Logania  watsoniana  watsoniana  de  Niceville;  Evans,  1932:  213. 

Logania  watsoniana  subfasciata  Tytler;  Evans,  1932:  213. 

Structurally  L.  watsoniana  is  closer  to  L.  distanti  than  to  L.  marmorata,  with  which,  probably  because  of  its 
crenulate  forewing,  it  has  sometimes  been  confused.  It  can  easily  be  separated  from  the  latter  by  its  legs, 
which  are  even  longer  than  those  of  L.  distanti  and  are  buff-brown  streaked  and  speckled  with  whitish. 


Fig.  48     Logania  watsoniana  de  Niceville;  Burma.  Male  genitalia. 


74  J.   N.  ELIOT 

In  the  wet  season  form,  subfasciata,  the  male  is  blackish  with  a  small  bluish  grey  discal  patch  in  spaces  3 
and  4  of  the  forewing,  just  extending  into  the  upper  part  of  space  2  and  the  base  of  space  5.  The  hindwing 
may  have  a  few  grey  scales.  The  female  has  a  more  extensive  pale  patch,  outwardly  white  and  inwardly  dark 
bluish  grey,  which  may  extend  to  the  wing  base  and  has  its  outer  edge  right-angled  at  vein  M3.  The 
underside  is  much  like  that  of  L.  marmorata. 

In  the  dry  season  form,  watsoniana,  the  pale  patch  on  the  forewing  of  the  male  extends  to  the  wing  base 
and  is  outwardly  white  and  angled  on  its  outer  edge,  and  the  hindwing  is  dusted  with  grey  below  vein  M\ .  In 
the  female  the  pale  areas  are  more  extensive  and  mostly  white,  and  on  the  underside  of  the  forewing  there 
is  a  large  white  area  beyond  and  below  the  cell. 

DISTRIBUTION.  Manipur;  Burma  as  far  south  as  Tavoy;  Thailand. 

Genus  LONTALIUS  gen.  n. 

Type-species:  Lontalius  eltus  sp.  n.  Gender  masculine.  The  generic  name  is  an  anagram  oi  Allotinus. 

The  genus  is  described  from  single  females  from  Pulo  Laut  and  Samar. 

Eyes  glabrous.  Antennae  similar  to  those  of  Paragerydus,  with  slender  shaft  and  gradually  incrassate 
club;  segments  number  about  55;  nudum  tapering  almost  to  base  of  shaft.  Labial  palpi  comparatively  short, 
resembling  those  of  Logania  rather  than  Allotinus,  with  third  segment  barely  longer  than  one-quarter  of 
second  segment.  Legs  (Fig.  49)  unlike  those  of  other  miletine  genera,  about  as  long  as  those  of  Allotinus, 
generally  thin,  but  with  tibiae  slightly  swollen;  long  first  segment  of  fore-tarsus  arched,  clothed  with  long 
hairs  in  place  of  normal  scales,  while  the  scales  on  remaining  tarsal  segments  much  longer  than  those  on 
femur  and  tibia  but  in  other  respects  of  normal  type.  Wings  crenulate,  with  a  particularly  long  tooth  on 
forewing  at  end  of  vein  M3.  Venation  (Fig.  49)  especially  distinguished  by  a  strong  humeral  vein  on 
hindwing  which  reaches  a  little  over  half-way  across  base  of  space  8.  The  single  species  is  moderately  large, 
brown  on  the  upperside  and  marked  on  the  underside  much  as  in  Allotinus  (Fabitaras)  fabius . 

This  monotypic  genus  appears  to  fall  between  Allotinus  and  Logania,  having  the  antennae,  pattern  and 
size  of  the  former,  while  the  crenulate  wings  and  short  labial  palpi  are  characteristic  of  the  latter,  but  it 
differs  from  both  in  the  peculiar  fore-tarsi  and  strong  humeral  vein. 


Fig.  49    Lontalius  eltus  treadawayi  subsp.  n.,  $;  Samar.  Left,  venation,  indicating  system  of  veins  and 
spaces  used  in  the  text;  centre,  mid-leg;  right,  fore-leg. 


THE  MILETINI  75 

Lontalius  eltus  sp.  n. 

(Figs  49,  venation  and  legs;  82  9 ;  108  9) 
The  characters  are  those  of  the  genus.  The  female  sex  only  is  known. 

Key  to  the  subspecies  of  L.  eltus 

1     Upperside  reddish  brown.  Underside  with  normal  lycaenid  markings  present  and  partly 

catenulate  on  hindwing eltus  eltus  (p.  75) 

Upperside  dull  brown  without  reddish  tinge.  Underside  with  lycaenid  markings  barely  de- 
veloped        eltus  treadawayi  (p.  75) 

Lontalius  eltus  eltus  subsp.  n. 

(Fig.  82  9) 

9  forewing  length  20-0  mm.  Upperside  reddish  brown;  cilia  pale  buff  but  dark  brown  at  vein  endings, 
broadly  so  on  forewing  and  narrowly  so  on  hindwing.  Underside  greyish  white,  freckled  with  chocolate- 
brown  specks  and  striae  which  are  more  or  less  coalesced  to  give  lycaenid-type  markings.  On  forewing 
these  comprise  a  series  of  broad  postdiscal  spots  in  spaces  4,  5  and  6,  a  broad  cell-end  bar  and  two  spots  in 
cell;  on  hindwing  a  complete  postdiscal  series  of  which  the  spots  in  spaces  2,  3  and  4  are  inwardly 
darker-edged  and  catenulate,  as  well  as  a  large  spot  mid-space  7  and  a  smaller  spot  in  inner  half  of  space  Ib. 
Cilia  as  on  upperside. 

Venation  differs  from  that  shown  in  Fig.  49  -  on  the  forewing  veins  Cuv  and  M3  are  connate,  on  the 
hindwing  the  humeral  vein  is  straight,  while  the  forewing  termen  is  more  strongly  dentate  at  vein  Cu2. 

Antennae  with  55  segments;  ringed  on  dorsal  side  of  shaft  with  dark  brown  and  whitish;  club  whitish  in 
basal  third,  remainder  dark  brown  except  for  unsealed  tip;  nudum  reddish  brown  tapering  gradually  to 
within  four  segments  of  base  of  shaft.  Labial  palpi  freckled  with  adpressed  dark  brown  and  whitish  scales. 
Legs  uniformly  pale  buff.  Body  brown  above  and  buff  below. 

MATERIAL  EXAMINED 

Holotype  9,  Borneo:  Pulo  Laut,  vi.1891  (W.  Doherty)  (BMNH). 

Unique.  The  specimen  is  labelled  /Type  [red]/Puro  Laut  Borneo  June  1891  Doherty/Borneo  Allotinus  n. 
sp.  9/Elwes  coll.  1915-207/Sp.  inc.  so  named  in  Elwes  coll. /Body  loose  and  fixed  7. v.  1948.  A.S.C./ 
Allotinus  eltus  Cbt.  9  H.T./.  Shortly  after  affixing  the  last  two  labels  Corbet  died,  and  the  name  eltus  was 
not  published. 

Lontalius  eltus  treadawayi  subsp.  n. 

(Figs  49,  venation  and  legs;  108  9) 

9  forewing  length  19-5  mm.  Upperside  dark  brown  without  a  reddish  tinge.  Cilia  mainly  worn  away,  but 
appear  to  be  similar  to  those  of  subsp.  eltus.  Underside  greyish  white  freckled  with  brown  as  in  subsp.  eltus, 
but  lycaenid  markings  not  apparent,  the  specks  being  only  slightly  denser  to  give  an  indication  of  postdiscal 
spots  in  spaces  4,  5  and  6  on  forewing  and  on  hindwing  in  spaces  6  and  7  as  well  as  a  suggestion  of  a  spot  in 
basal  half  of  space  7. 

MATERIAL  EXAMINED 

Holotype  9>  Philippines:  central  Samar,  Bagacay,  900  ft,  ll.viii.1979  (G.  C.  Treadaway)  (coll. 
Treadaway).  Regrettably  the  specimen  is  worn  and  in  tattered  condition,  with  the  antennae  broken  after  38 
segments  and  one  middle  leg  and  both  hind  legs  missing. 

< 

Genus  MILETUS  Hiibner 

Miletus  Hiibner,  1819:  71.  Type-species:  Papilla  symethus  Cramer,  1779,  by  designation  of  Doubleday, 

Westwood  &  Hewitson,  1852:  502.  Gender  masculine. 

Symetha  Horsfield,  1828:  59.  Type-species:  Symetha pandu  Horsfield  1828,  by  monotypy. 
Gerydus  Boisduval,  1836:  pi.  23,  fig.  2.  Type-species:  Papilio  symethus  Cramer,  1779,  by  original 

designation. 
Archaeogerydus  Fruhstorfer,  1916:  816.  Type-species:  Gerydus  croton  Doherty,  1889,  by  designation  of 

Hemming,  1960:  9. 


76 


J.   N.   ELIOT 


The  genus  is  instantly  recognisable  by  three  external  characters.  The  first  segment  of  the  tarsi  is  long, 
flattened  and  blade-like.  There  is  no  humeral  vein  on  the  hindwing.  The  pattern  on  the  underside 
comprises  normal  catenulate  lycaenid  markings  on  a  ground  which  is  not  striated  or  freckled.  The  male 
genitalia  are  typical  of  the  tribe,  with  the  arms  of  the  juxta  particularly  strongly  conjoined  above  the 
phallus.  The  twin  abdominal  hair  tufts  on  the  sternum  of  the  eighth  abdominal  segment  are  usually 
permanently  extruded. 

The  genus  is  distributed  from  India  to  south  China  and  through  the  Malay  Archipelago  and  Philippines 
to  New  Guinea.  It  was  revised  by  Eliot  (1961),  and  I  have  no  reason  to  alter  the  arrangement  proposed 
therein  with  the  exception  of  a  few  amendments  detailed  below.  However,  as  there  are  now  a  few 
additional  species,  I  have  thought  it  best  to  include  herein  a  fresh  key  to  the  species.  Opportunity  has  been 
taken  to  figure  the  new  species  as  well  as  a  few  previously  unfigured  taxa,  viz.  M.  nymphis  eneus  (Fig.  85, 
Cf),  M.  cellarius  (Figs  86,  cf ,  87,  $),  M.  heradeion  (Figs  94,  96,  cf ,  95  $),  M.  mallus  mallus  (Fig.  83,  Cf) 
and  M.  celinus  (Fig.  104,  cf). 

Key  to  the  species  of  Miletus 

1  cf  valva  with  distal  half  not  tapering  distad,  dorsal  edge  not  narrowly  folded  inwards,  cf 

forewing  vein  M3  unswollen  or  basally  swollen  and  clothed  with  specialised  scales 2 

Cf  valva  more  or  less  tapering  distad,  dorsal  edge  narrowly  folded  inwards  (Fig.  50D).  cf 
forewing  with  vein  M3  always  swollen  and  clothed  with  specialised  scales  (symethus-group) 

2  cf  distal  third  of  valva  not  trough-shaped .  cf  forewing  vein  M3  swollen  or  unswollen 

Cf  distal  third  of  valva  more  or  less  rectangular,  with  edges  curved  inwards  to  form  a  U-shaped 

trough  (Fig.  50C) .  cf  forewing  vein  M3  swollen  (boisduvali-group) 

3  cf  distal  third  of  valva  subspatulate,  with  dorsal  margin  less  convex  than  ventral  margin,  and 

with  a  terminal  hook  at  apex  (Fig.  50A)  (cMnens/s-group) 

Cf  distal  third  of  valva  spatulate,  with  terminal  hook  very  small  and  in  centre  line  (Fig.  SOB) 
(zinclcenii-group) 

4  cf  forewing  vein  M3  not  or  only  weakly  swollen ,  partly  covered  with  normal  scales 

Cf  forewing  vein  M3  strongly  swollen  and  clothed  with  specialised  scales M.  chinensis 

5  Cf  $  upperside  of  forewing  with  white  to  whitish  markings,  which  may  be  heavily  sullied  in  wet 

season  forms,  comprising  a  patch  beyond  end-cell  and  smaller  and  usually  separate  spots  in 

spaces  2  and  Ib.  cf  forewing  vein  M3  not  swollen 

Cf  9  upperside  of  forewing  not  so  marked .  cf  forewing  vein  M3  weakly  swollen  or  unswollen .... 

6  Larger,  forewing  20-24  mm.  Underside  of  hindwing  rather  dark,  variegated  brown,  usually 

becoming  more  or  less  blackish  on  disc,  with  reddish  brown  markings.  Underside  of  forewing 

without  a  whitish  streak  above  dorsum M.  croton 

Smaller,  forewing  14-22  mm.  Underside  of  hindwing  greyish  to  buff-brown,  with  markings 
only  slightly  more  reddish  than  ground  colour.  Underside  of  forewing  with  a  more  or  less 

developed  whitish  streak  above  dorsum M.  mallus 

1     Cf  $  upperside  of  forewing  with  a  continuous  and  more  or  less  even  white  band        M.  nymphis 
Cf  $  upperside  of  forewing  usually  unmarked  brown,  but  markings  similar  to  those  of  M. 
croton  and  M.  mallus  are  occasionally  faintly  discernible  (Fig.  84),  most  often  in  $ 

M.  gaesa 


14 
3 

11 


C  D 

Fig.  50    Valvae  of  species-groups  of  Miletus.  A,  M.  chinensis  C.  Felder;  B,  M.  zinckenii  C.  &  R.  Felder; 
C,  M.  boisduvali  Moore;  D,  M.  symethus  (Cramer). 


THE  MILETINI  77 

8  cf  vein  M3  strongly  swollen  and  clothed  with  specialised  scales 9 

-  cf  vein  M3  unswollen  and  clothed  with  normal  scales 10 

9  Cf  $  upperside  of  forewing  with  white  band  outwardly  more  or  less  straight  and  oblique  from 

vein  A  i  to  vein  Af3,  thence  curved  basad.  $  hindwing  strongly  produced  at  vein  M3 

M.  gopara 

-  Cf  $  upperside  of  forewing  with  white  band  outwardly  angled  at  vein  Cui ,  below  which  it  is  at 

right  angles  to  dorsum.  9  hindwing  normal,  barely  toothed  at  vein  M3 M.  zinckenii 

10  cf  $  upperside  of  hindwing  unicolorous  brown M.  valeus 

-  Cf  $  upperside  of  hindwing  mostly  white M.  gaetulus 

11  Cf  upperside  of  forewing  with  white  or  whitish  markings,  except  in  smokey  brown  M.  drucei 

sometimes 12 

-  cf  upperside  of  forewing  brown  with  a  sepia  tinge  and  unmarked M.  boisduvali 

12  cf  $  upperside  of  forewing  with  white  band  not  extending  basad  of  outer  quarter  of  cell 13 

-  Cf  $  upperside  of  forewing  with  broad  white  band  filling  outer  half  of  cell. 

Underside  markings  very  clearly  defined  (Figs  86, 87) M.  cellarius 

13  cf  very  variable;  on  upperside  of  forewing  white  markings  may  be  absent,  or  form  a  diffuse 

whitish  spot  beyond  end-cell  in  spaces  3-5,  or  may  be  clear  white  and  extend  into  spaces  2 
and  Ib  commencing  at  base  of  space  2. 

$  usually  with  a  more  or  less  circular  white  patch  beyond  end-cell,  but  a  white  band  similar 
to  that  of  well-marked  males  may  be  present.  Philippines  and  North  Borneo         M.  drucei 

-  cf  less  variable;  white  markings,  at  their  most  extensive,  do  not  reach  base  of  space  2 

M.  biggsii 

14  cf  outer  half  of  valva  tapering  more  or  less  evenly  and  ending  in  a  blunt  point  or  narrowly 

truncate  (Figs  50D,  51) 15 

-  Cf  outer  half  of  valva  tapering  unevenly,  apex  broadly  truncate  or  broadly  rounded  (Figs  52, 

53) 24 

15  Underside  of  hindwing  with  postdiscal  markings  in  spaces  4  and  5  equidistant  from  end-cell  and 

termen ,  or  very  nearly  so 16 

-  Underside  of  hindwing  with  postdiscal  markings  in  spaces  4  and  5  much  closer  to  termen  than 

end-cell 23 

16  Forewing  apex  without  a  protruding  point 17 

-  Forewing  apex  produced  to  a  short  point,  terrhen  concave  in  space  6  (Figs  102, 103) 

M.  takanatnii 

17  Upperside  of  hindwing  with  discocellular  veins  not  darkened ,  without  bluish  grey  scaling 18 

-  Upperside  of  hindwing  with  discocellular  veins  more  or  less  darkened,  with  bluish  grey  scaling 

(except  in  some  Philippine  subspecies)  (Figs  88-9 1 ) M.  symethus 

18  Upperside  forewing  base  and  all  hindwing  brown  (but  hindwing  may  be  sullied  with  white 

scales  in  dry  season  M.  ancon) 19 

-  Upperside  forewing  base  white  and  hindwing  almost  all  pure  white 22 

19  Upperside  of  forewing  with  white  band  (if  present)  constricted  and  may  be  completely  divided 

below  basal  half  of  vein  Cui 20 

-  Upperside  of  forewing  with  white  band  not  constricted  below  vein  Cu\ 21 

20  Upperside  blackish  brown.  Forewing  with  a  broad  white  band  from  vein  Sc  to  dorsum  which  is 

constricted  and  may  be  narrowly  divided  below  vein  Cu\,  at  least  one-third  of  space  2 
white M.  ancon 

-  Upperside  more  reddish  brown.  Forewing  with  white  markings  much  reduced  (absent  in  cf  of 

nominate  subspecies),  with  lower  part  of  band  widely  separated  from  upper  part;  in  space  2 
at  most  a  round  white  spot M.  archilochus 

21  Underside  of  hindwing  with  markings  in  cell  and  spaces  3  and  4  darker  than  other  markings. 

Smaller,  forewing  17-18  mm M.  gallus 

-  Underside  of  hindwing  with  all  markings  of  more  or  less  same  intensity.  Larger,  forewing  19-23 

mm M.  heracleion 

22  Underside  of  forewing  with  a  black  discal  area  which  more  or  less  completely  blacks  out  cell 

markings M.  gigantes 

-  Underside  of  forewing  with  cell  markings  not  blacked  out M.  atimonicus 

23  Forewing  termen  and  apex  normal M.  leos 

-  Forewing  apex  produced  to  a  short  point,  termen  strongly  convex  (Fig.  104) M.  celinus 

24  cf  valva  with  apex  broadly  truncate,  slightly  concave  (Fig.  52) M.  melanion 

-  Cf  valva  with  apex  broadly  rounded  (Fig.  53) M.  bazttanus 


78  J.   N.  ELIOT 

Miletus  mallus  (Fruhstorfer) 

(Fig.  83  cT) 
Gerydus  croton  mallus  Fruhstorfer,  1913:  310. 

In  my  original  analysis  of  Miletus  (Eliot,  1961)  I  treated  M.  mallus  as  a  good  species,  but  later  (1967:  72)  I 
suggested  that  it  was  conspecific  with,  and  a  subspecies  of,  M.  gaesa.  At  the  same  time  I  erroneously 
recorded  a  montane  form  or  microsubspecies  of  M.  gaesa  (Fig.  84)  from  the  mountains  of  the  Malay 
Peninsula  as  M.  gaesa  mallus.  I  now  consider  that  M.  mallus  and  M.  gaesa  should  be  treated  as  distinct 
allopatric  species. 

Until  recently  I  was  only  able  to  examine  four  specimens  of  M.  mallus  from  Vietnam,  namely  the  male 
holotype  of  mallus  from  south  Annam,  which  was  taken  in  the  dry  season,  and  the  male  holotype  and  two 
females  of  gethusus,  which  were  taken  in  Tonkin  in  the  wet  season.  I  presumed,  mistakenly,  that  these 
represented  seasonal  forms  of  a  single  taxon.  I  have  now  been  able  to  examine  a  series  of  mallus  taken  by 
Bedford  Russell  at  Dalat  in  south  Vietnam  during  the  wet  season.  The  males  differ  hardly  at  all  from  the 
holotype  of  mallus,  but  differ  from  gethusus  in  several  particulars,  notably  in  having  quite  well-developed 
whitish  postdiscal  markings  on  the  upperside  of  the  forewing  and  a  blackish  area  in  and  beneath  the  cell  on 
the  underside.  In  consequence  I  consider  that  gethusus  should  be  reinstated  as  a  valid  subspecies  of  M. 
mallus  flying  in  north  Vietnam. 

Miletus  symethus  (Cramer) 
(Figs  88  $;  89,  90  cf;  91$) 
Papilio  symethus  Cramer,  1779:  84. 

In  my  analysis  (1961)  I  suggested  that  the  dark  subspecies  philopator  from  Mindoro  probably  occurred  in 
Luzon  and  other  Philippine  islands.  I  have  now  seen  females  from  Luzon  and  both  sexes  from  Marinduque, 
which  bear  little  resemblance  to  philopator  and  constitute  a  new  subspecies. 

Miletus  symethus  phantus  subsp.  n. 

(Fig.  88$) 

Cf  broadly  similar  to  subsp.  edonus  from  Palawan,  from  which  it  differs  by  the  reduction  of  the  greyish  blue 
scaling  on  the  hindwing,  so  that  the  lower  half  of  the  wing  looks  greyish  brown. 

$  differs  from  subsp.  edonus  in  the  same  way  as  the  male,  namely  in  the  considerably  browner  hindwing, 
which  usually  has  a  white  streak  in  the  basal  three-quarters  of  spaces  4  and  5  and  in  the  extreme  end  of  the 
cell. 

The  subspecies  approaches  subsp.  hierophantes  (Fig.  89)  from  the  Sulu  Is  and,  in  a  form  with  a  much 
darker  underside  (Figs  90,  91),  from  Mindanao,  in  which  the  usual  bluish  grey  scaling  on  the  hindwing  is 
entirely  absent,  so  that  the  non-white  area  of  the  wing  is  dark  brown. 

MATERIAL  EXAMINED 

Holotype  cf ,  Philippines:  Marinduque,  iv.1980  (coll.  Treadaway). 

Paratypes.  Philippines:  1  $  (allotype),  Marinduque,  vii.1979  (ex  coll.  Takanami)  (BMNH);  1  $ ,  data  as 
holotype  (coll.  Treadaway). 

Excluded  from  type-series.  Philippines:  2  $ ,  Luzon,  Bicol  National  Park,  28-29. viii.l 980  (Y.  Takanami) 
(BMNH).  These  differ  from  Marinduque  females  by  a  larger  white  area  on  the  underside  of  the  forewing; 
in  addition,  in  one  example  (Fig.  88)  the  white  streak  on  the  hindwing  is  obsolescent. 


Miletus  ?  symethus  solitarius  Okubo 

Miletus  ancon  solitaria  Okubo,  1983:  176,  figs  27,  28  cf .  Holotype  cf ,  WEST  MALAYSIA:  Tioman  I.  (coll. 
Okubo). 

I  have  not  seen  the  unique  holotype  from  Tioman,  an  island  some  30  miles  off  the  east  coast  of  the  Malay 
Peninsula.  Okubo's  figures  show  a  butterfly  in  which  the  white  markings  on  the  forewing  comprise  a 
triangular  patch  beyond  the  end  of  the  cell  which  is  separated  by  a  broad  dark  bar  from  a  smaller,  elongated 


THE  MILETINI  79 

spot  in  space  2,  a  larger  more  or  less  contiguous  spot  in  space  Ib  and  a  narrow  white  streak  in  space  la  above 
the  centre  of  the  dorsum.  I  do  not  think  it  can  be  a  subspecies  of  M.  ancon  since  it  possesses  several 
discordant  characters.  It  is  smaller  than  any  ancon  I  have  seen,  the  forewing  length  being  only  18-0  mm;  the 
base  of  the  forewing  and  all  the  hindwing  below  vein  MI  are  paler;  the  discocellular  veins  of  the  hindwing 
are  darkened;  and  on  the  underside  of  the  forewing  the  markings  in  the  cell  are  not  blacked  out.  In  fact,  the 
butterfly  accords  much  better  with  M.  symethus  than  with  any  other  species,  and  broadly  resembles  a 
frequent  male  variety  of  the  nominate  subspecies  in  Java,  in  which  the  white  markings  are  similarly 
reduced. 

The  only  two  males  of  M.  symethus  from  Tioman  which  I  have  examined  differ  from  subsp.  petronius 
from  the  nearby  mainland  in  having  the  basal  area  of  the  forewing  darker  and  more  extensive,  so  that  the 
white  markings  are  reduced  though  not  divided.  On  these  grounds  the  local  Tioman  race  appears  to  be  a 
good  subspecies.  I  suspect  that  the  holotype  ofsolitarius  is  just  a  variety  of  this  race,  in  which  the  tendency 
for  a  darker  and  enlarged  basal  area  is  much  increased.  I  therefore  treat  the  name  solitarius  conditionally  as 
that  of  the  symethus  subspecies  flying  in  Tioman  I. 


Miletus  archilochus  siamensis  (Godfrey) 

Gerydus  ancon  siamensis  Godfrey,  1916:  134.  Holotype  cf ,  THAILAND:  east  (BMNH)  [examined]. 
Miletus  archilochus  (Fruhstorfer);  Lewis,  1974:  pi.  178,  fig.  29  cf . 

In  my  analysis  of  1961 1  retained  siamensis  as  a  subspecies  of  M.  ancon,  as  I  thought  that  the  two  taxa  were 
allopatric.  Since  then  I  have  seen  examples  of  nominate  ancon  from  throughout  Thailand  apparently  flying 
sympatrically  with  siamensis.  So  wide  an  overlap  argues  against  conspecificity,  and  I  now  think  that 
siamensis  must  be  a  subspecies  of  the  sibling  species  M.  archilochus.  In  siamensis  the  rather  sullied  white 
markings  on  the  forewing  decrease  in  extent  from  west  to  east.  These  markings  are  absent  in  the  male  of 
nominate  archilochus  and  present,  but  obscure,  in  the  female.  I  have  seen  no  examples  from  the  territory 
between  Thailand  and  Tonkin  (type-locality  of  archilochus},  and  I  anticipate  that  these  will  prove  to  be 
intermediate  and  show  that  variation  is  clinal. 


Miletus  gigantes  (de  Niceville) 
Gerydus  gigantes  de  Niceville,  1894:  23. 

In  my  analysis  (1961)  I  considered  this  species  to  be  a  subspecies  of  M.  ancon  because  of  the  absence  of  any 
known  overlap  in  their  distribution.  Later  (1978:  237)  I  restored  gigantes  to  specific  status.  The  phenotype, 
with  the  upperside  mainly  white,  differs  greatly  from  that  of  M.  ancon  and,  in  addition,  there  is  an 
apparently  constant  difference  in  the  male  genitalia.  In  continental  and  Bornean  ancon  there  is  only  a 
single  spine-like  cornutus  in  the  phallus,  whereas  in  gigantes  there  are  two  spines  as  in  most  other  species  of 
the  genus. 


Miletus  atimonicus  Murayama  &  Okamura  stat.  n. 

(Figs  51,  cf  genitalia;  92  cf ;  93  $) 

Miletus  sumethus  [sic]  atimonicus  Murayama  &  Okamura,  1973:  23,  figs  45,  46  'cf'  recte  9-  Holotype  'cf' 
recte  $,  PHILIPPINES:  Luzon  (coll.  Murayama)  [examined]. 

Hitherto  only  known  from  the  female  holotype,  which  the  authors  mistook  for  a  male  of  a  M.  symethus 
subspecies.  Recently  I  have  been  sent  a  male  from  Negros  and  a  female  from  Luzon  by  Mr  Yusuke 
Takanami.  Both  sexes  bear  a  superficial  resemblance  on  both  surfaces  to  M.  gaetulus  (de  Niceville,  1894), 
and  on  the  upperside  to  M.  gigantes.  In  the  male  the  basal  third  of  vein  M3  of  the  forewing  is  swollen  and 
clothed  with  specialised  scales;  this  character  immediately  separates  it  from  M.  gaetulus  wherein  it  is 
absent.  It  also  differs  from  the  latter  on  the  underside  by  a  browner  ground  colour  and  more  reduced 
markings  in  the  forewing  cell.  The  underside  is  not  at  all  like  that  of  M.  gigantes,  lacking  the  prominent 
blackish  discal  areas  found  on  the  forewing  of  that  species. 

The  male  genitalia  indicate  that  M.  atimonicus  belongs  to  the  symethus-group.  However,  the  valva  is 
truncate  just  before  the  apex  and  shows  a  slight  approach  to  the  valva  of  M.  melanion.  The  phallus,  with  a 
single  spine-like  cornutus,  recalls  that  organ  in  M.  ancon. 


80 


J.  N.  ELIOT 


Fig.  51     Miletus  atimonicus  Murayama  &  Okamura;  Negros.  Male  genitalia. 

Miletus  heracleion  (Doherty) 

(Figs  94,  96  cf;  95$) 
Gerydus  heracleion  Doherty,  1891ft:  36. 

This  species  has  not  been  recorded  from  Sumatra,  but  there  is  in  BMNH  a  male  (West  Sumatra,  Lebong 
Tandai,  March  1923,  C.  J.  Brooks)  which  is  somewhat  larger  but  otherwise  hardly  differs  from  the 
nominate  subspecies.  There  is  also  a  male  from  Sarawak  (Bidi,  February  1909,  C.  J.  Brooks)  (Fig.  94) 
which  is  similar  to  the  Sumatran  male  except  that  on  the  underside  of  the  forewing  the  white  band  crossed 
by  dark-dusted  veins  is  little  more  than  half  as  wide.  It,  and  a  female  from  Pulo  Laut  (Fig.  95)  previously 
recorded  as  subsp.  arion  (Fig.  96)  (Eliot,  1961:  171),  but  which  has  the  white  band  on  the  forewing 
considerably  narrower  on  the  upperside  but  wider  on  the  underside  than  in  that  subspecies,  possibly 
represent  a  further  minor  subspecies,  but  they  are  provisionally  placed  under  subsp.  heracleion. 

Miletus  melanionC.  &  R.  Felder 
(Figs  52,  cf  genitalia;  97,  98  C?) 
Miletus  melanion  C.  &  R.  Felder,  1865:  284. 

It  has  been  found  that  melanion,  as  treated  by  all  authors  up  to  and  including  Eliot  (1961),  comprises  two 
superficially  similar  species  with  different  male  genitalia,  namely  M.  melanion  and  M.  bazilanus  (Fruhstor- 
fer,  1913).  The  former  occurs  throughout  the  Philippines  (excluding  Palawan)  at  least  as  far  south  as 
Mindanao,  whilst  the  latter  is  at  present  known  only  from  Mindanao  and  Bazilan.  This  distribution 
suggests  that  the  two  species  evolved  in  the  northern  and  southern  island  groups  respectively,  and  that 
melanion  was  later  able  to  spread  south  into  Mindanao,  where  to-day  it  is  a  common  species,  without 
interbreeding  with  the  endemic  bazilanus. 

M.  melanion  exhibits  a  good  deal  of  individual  variation,  but  broadly  the  males  can  be  broken  down  into 
two  dimorphs,  which  at  first  sight  look  as  though  they  should  represent  distinct  species.  The  male  genitalia, 
however,  reveal  no  significant  differences,  and  the  females  cannot  be  separated  into  two  groups.  In  the 
typical  male  (Fig.  97),  figured  by  the  Felders  (1865:  pi.  35,  figs  32,  33),  the  brown  forewing  has  a  short, 
more  or  less  sullied  white  subtornal  streak  in  space  Ib  and  the  swollen  portion  of  vein  M3  is  not  surrounded 
by  a  white  area,  although  a  thin  band  of  whitish  scales  below  and  a  sullied  white  spot  above  the  swelling  may 
be  present.  On  the  underside  of  the  forewing  there  are  sullied  white  subtornal  spots  in  spaces  Ib  and  2  and  a 
sullied  whitish  streak  beyond  the  cell.  The  dimorph,  cf-f.  albiguttatus  f.  n.  (fig.  98)  (holotype  cf ,  Luzon, 
Lorquin,  ex  Felder  coll.  (BMNH))  is  generally  smaller  and  more  blackish  brown  above;  there  is  an 
additional  whitish  subtornal  spot  in  space  2  and  a  prominent,  usually  clear  white,  more  or  less  triangular 
spot  surrounding  the  swollen  portion  of  vein  M3,  with  its  base  extending  from  mid-space  3  to  the  cell  apex. 


THE  MILETINI 


81 


On  the  underside  of  the  forewing  there  is  a  broad  white  area  crossed  by  dark-dusted  veins,  extending  from 
the  dorsum  to  vein  MI. 

The  female  generally  resembles  cf-f.  albiguttatus,  but  the  white  spots  are  usually  larger.  I  do  not  think 
they  are  ever  conjoined  into  a  continuous  white  band,  such  as  occurs  frequently  in  M.  bazilanus. 

In  the  male  genitalia  the  valva  is  slightly  upturned  and  truncate  before  the  apex  and  its  outer  edge  is 
slightly  concave. 

Key  to  the  subspecies  of  M.  melanion 

1  cf  dimorphic.  $  upperside  of  forewing  with  white  markings  comprising  a  sullied  streak  in  space 
la,  a  double  subtornal  spot  in  space  Ib,  an  overlapping  spot  in  space  2  separated  by  a  dark  bar 
from  a  much  larger  spot  beyond  the  cell  which  extends  above  vein  R5,  often  as  far  as  vein  Sc 

melanion  melanion  (p.  81) 

Cf  monomorphic,  apparently  only  f.  melanion  being  present.  $  upperside  of  forewing  with 
subtornal  white  markings  normally  reduced  to  an  often  sullied  white  streak  in  upper  part  of 
space  Ib,  with  the  white  spot  beyond  the  cell  smaller,  not  extending  above  vein  R5 

melanion  euphranor  (p.  82) 

Miletus  melanion  melanion  C.  &  R.  Felder 
(Figs  52  cf  genitalia;  97,  98  cf ) 

Miletus  melanion  C.  &  R.  Felder,  [1865]:  284,  pi.  35,  figs  32,  33  cf .  LECTOTYPE  cf ,  PHILIPPINES:  Luzon 

(BMNH),  here  designated  [examined]. 

Gerydus  melanion  (C.  &  R.  Felder);  Semper,  1889:  161,  pi.  31,  figs  13,  14  $. 
Gerydus  melanion  melanion  (Felder);  Fruhstorfer,  1913:  246;  1916:  822,  pi.  14d  cf . 
Miletus  melanion  melanion  C.  &  R.  Felder;  Eliot,  1961:  175,  partim. 

The  characters  of  the  subspecies  are  given  in  the  key. 

Judging  by  material  in  BMNH,  in  Luzon  f.  melanion  nearly  always  has  a  small,  often  sullied,  white  spot 
above  the  swollen  portion  of  vein  M3.  In  the  southern  group  of  islands  this  white  spot  is  seldom  present, 
nearly  all  males  resembling  Fig.  97.  It  seems  that  fully  marked  f.  albiguttatus  is  rare  in  Luzon,  and  many 
examples  are  intermediate  to  f.  melanion;  elsewhere  the  two  forms  remain  more  sharply  distinct. 

In  BMNH  there  are  three  males  from  the  Felders'  type-series,  one  of  which  is  f.  albiguttatus  already 
designated  as  holotype.  The  other  two  are  f.  melanion  with  a  sullied  white  spot  above  vein  M3. 1  designate 
as  lectotype  the  one  which  most  nearly  resembles  the  example  figured  by  the  Felders  and  has  the  white  spot 
very  small  and  sullied;  it  is  labelled  /80/Luzon  Lorquin  [round  blue]/Melanion  n. /Felder  Colin,  [round 
white]/Rothschild  Bequest  B.M.  1939-1/. 

DISTRIBUTION.  Throughout  the  Philippines,  excluding  Mindoro,  Palawan  and  the  Sulu  Is. 


Fig.  52     Miletus  melanion  melanion  cf-f.  albiguttatus  f.  n.;  Negros.  Male  genitalia.  Left,  above,  latero- 
dorsal  view  of  interior  of  left  valva  and,  below,  ventral  view  of  both  valvae. 


82 


J.  N.  ELIOT 


Miletus  melanion  euphranor  (Fruhstorfer) 


Gerydus  melanion  euphranor  Fruhstorfer,  1914:  60;  1916:  822.  LECTOTYPE  cf ,  PHILIPPINES:  Mindoro 

(BMNH),  here  designated  [examined]. 
Miletus  melanion  melanion  C.  &  R.  Felder;  Eliot,  1961:  175,  partim,  fig.  24  cf  genitalia. 

The  characters  of  this  dark  subspecies  are  given  in  the  key. 

In  BMNH  there  are  two  male  and  one  female  syntypes.  I  designate  as  lectotype  one  of  the  males, 
labelled:  /Syntype  [blue]/Baco  Dist.  Mindoro  6.5.09/Adams  Bequest  B.M.  1912-399/;  its  genitalia  were 
figured  by  Eliot  (1961:  fig.  24).  The  other  male  and  female  are  paralectotypes. 

DISTRIBUTION.  Mindoro.  Also  Leyte,  whence  some  examples  in  BMNH  tend  to  be  even  darker  than  those 
from  Mindoro  whilst  others  (Fig.  97)  are  inseparable  from  Luzon  examples;  it  might  have  been  expected 
that  subsp.  melanion  alone  would  have  occurred  there. 

Miletus  bazilanus  (Fruhstorfer)  stat.  n. 
(Figs  53  cf ,  genitalia;  99  cf ;  100, 101  $) 

Gerydus  melanion  bazilanus  Fruhstorfer,  1913:  246;  1916:  822.  LECTOTYPE  cf ,  PHILIPPINES:  Bazilan 

(BMNH),  here  designated  [examined]. 
Gerydus  melanion  vitelianus  Fruhstorfer,  1913:  246;  1916:  822.  LECTOTYPE  $,  PHILIPPINES:  Mindanao 

(BMNH),  here  designated  [examined].  Syn.  n. 
Miletus  melanion  vitelianus  (Fruhstorfer)  Eliot,  1961:  175,  partim. 
Miletus  melanion  bazilanus  (Fruhstorfer)  Eliot,  1961: 175. 

The  male  is  very  similar  to  sympatric  M.  melanion  f.  melanion,  but  on  the  upperside  of  the  forewing  the 
white  streak  in  space  Ib  is  usually  more  prominent  and  about  3-5  mm  long;  the  underside  is  slightly  darker 
and  the  postdiscal  markings  are  wider.  However,  to  be  absolutely  certain  of  identification  some  males  may 
need  dissection. 

The  females  generally  have  the  white  markings  more  extensive  than  M.  melanion,  and  in  typical 
examples  (Fig.  100)  there  is  a  continuous  white  band  on  the  forewing  extending  from  the  dorsum  to  vein  Sc. 
However,  some  examples  (Fig.  101)  resemble  normal  melanion  females,  except  by  a  slightly  darker 
underside  with  wider  postdiscal  markings,  and  these  are  connected  by  intermediates  to  the  typical  form. 

The  male  genitalia  differ  from  those  of  M.  melanion  in  three  respects.  The  valva  is  distally  broader,  with 
the  tip  rounded;  the  paired,  spine-like  cornuti  in  the  phallus  are  more  than  twice  as  large;  the  unci  have  a 
more  elongated  tip. 

I  designate  as  lectotype  of  bazilanus  a  male  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer  orange]/ 


Fig.  53     Miletus  bazilanus  (Fruhstorfer);  Mindanao.  Male  genitalia.  Left,  above,  latero-dorsal  view  of 
interior  of  left  valva  and,  below,  ventral  view  of  both  valvae. 


THE  MILETINI 


83 


Philippinen  Bazilan  II-III  98  Doherty  ex  coll.  Fruhstorfer/Fruhstorfer  Coll.  B.M.  1933-131/melanion 
bazilanus  Frhst.  [in  Fruhstorfer's  hand]/. 

I  designate  as  lectotype  of  vitelianus  a  female  in  BMNH  labelled  /Type  [red]/Type  [Fruhstorfer 
orangej/Fruhstorfer  Coll.  B.M.  1933-131/melanion  vitelianus  Frhst.  [in  Fruhstorfer's  hand]/;  it  has  vein 
Cui  dark-dusted  across  the  forewing  white  band.  There  are  no  males  from  Mindanao  ex  Fruhstorfer  coll.  in 
BMNH,  but  his  description  of  the  male  accords  reasonably  well  with  bazilanus. 

DISTRIBUTION.  Mindanao;  Bazilan. 

Miletus  takanamiisp.  n. 

(Figs  54,  cf  genitalia;  102  cf ;  103  £) 

Cf  forewing  length  19-0  mm.  Body  and  labial  palpi  dark  brown.  Legs  buff-brown.  Wing  shape  similar  to 
that  of  the  Celebesian  M.  celinus  (Fig.  104),  with  the  forewing  concave  in  space  6  below  a  pointed  apex,  and 
with  the  basal  quarter  of  vein  M3  swollen  and  clothed  with  the  usual  minute  wedge-shaped  specialised 
scales.  Upperside  blackish  brown ;  forewing  with  a  white  patch  3-0  mm  wide  surrounding  swollen  portion  of 
vein  M3,  extending  from  mid-space  2  across  spaces  3  and  4  and  into  space  5  as  a  narrow  projection,  and  a 
white  streak  5-5  mm  long  and  1-0  mm  wide  just  below  centre  of  vein  Cu2.  Underside  paler  brown  becoming 
reddish  near  middle  of  forewing  termen;  usual  markings  reddish  brown,  outlined  by  black  lines  except  on 
outer  side  of  postdiscal  series  of  hindwing.  Forewing  with  an  irregular  white  area  extending  from  just  above 
dorsum  to  vein  R5,  5-0  mm  wide  in  space  Ib,  increasing  to  6-5  mm  in  space  2,  then  decreasing  to  3-5  mm  in 
space  6.  Genitalia  with  valva  tapering  to  a  rounded  point,  as  in  most  species  of  the  symethus-group,  phallus 
short,  much  as  in  M.  melanion  and  M.  bazilanus,  but  lacking  cornuti. 

9  forewing  18-0  mm.  Forewing  termen  more  convex  than  in  male,  hindwing  slightly  dentate  at  vein  M3, 
as  usual  in  the  genus.  Upperside  blackish  brown;  forewing  with  an  irregular,  oblique  white  band  running 
from  vein  Al,  where  it  is  5-0  mm  wide,  to  vein  /?1?  where  it  is  2-0  mm  wide,  and  entering  outer  part  of  cell; 
veins  Cui  and  Cu2  are  dark-dusted  where  they  cross  this  band.  Underside  generally  similar  to  male,  but  on 
hindwing  lower  half  of  area  between  discal  and  postdiscal  spots  dusted  with  black  scales. 

MATERIAL  EXAMINED 

Holotype  cf ,  Philippines:  Mindanao,  Tandag,  Surigao,  1981-1982  (ex  coll.  Takanami)  (BMNH). 
Paratype.  1  $  (allotype),  ii.1982,  data  otherwise  as  holotype  (BMNH). 

Apart  from  its  distinctive  wing  shape  the  male  most  nearly  resembles  M.  melanion  f.  albiguttatus,  from 
which  it  differs  in  having  a  larger  white  patch  astride  the  swollen  portion  of  vein  M3  and  in  lacking  a  small 
white  subtornal  spot  in  space  2  and,  on  the  underside,  in  a  darker  and  more  irregular  ground  colour  and 


C 


P 


Fig.  54     Miletus  takanamii  sp.  n. ;  Mindanao.  Male  genitalia.  Lower  left,  latero-dorsal  view  of  interior  of 
left  valva. 


84  J-  N.  ELIOT 

wider  white  area  on  the  forewing.  The  female  is  much  like  that  sex  of  M.  bazilanus,  but  on  the  underside 
the  ground  colour  is  much  darker  and  the  white  central  area  of  the  forewing  is  wider. 
The  species  is  named  after  Mr  Yusuke  Takanami,  who  generously  presented  the  types  to  the  BMNH. 


Genus  MEGALOPALPUS  Rober 

Megalopalpus  Rober,  1886:  51.  Type-species:  Megalopalpus  simplex  Rober,  1886:  51,  pi.  4,  fig.  4,  by 
original  designation.  Gender  masculine.  [See  also  Opinion  566, 1959.] 

This  genus  appears  to  be  closely  allied  to  Miletus,  of  which  it  should  perhaps  be  treated  as  a  subgenus, 
having  similar  long  legs  with  the  foretarsi  flattened  and  blade-like,  and  the  undersurface  of  the  wings 
similarly  patterned  with  catenulate  markings,  although  these  are  sometimes  obsolete.  The  only  significant 
difference  lies  in  the  presence  of  a  strongly  developed  humeral  vein  in  Megalopalpus.  Males  also  lack 
secondary  sexual  characters,  but  this  condition  is  matched  in  some  species  belonging  to  the  Oriental 
genera.  The  male  genitalia  are  of  the  normal  pattern  for  the  tribe,  but  with  some  distinctive  characters:  the 
uncus/tegumen  plates  are  triangular  and  bear  a  broad,  lobe-like  process  directed  ventrad,  and  the  brachia 
are  curved  (see  illustrations  in  Bethune-Baker,  1914;  Stempffer,  1967;  Eliot,  1973). 

The  genus  is  restricted  to  the  forested  regions  of  West  Africa,  from  Liberia  south  to  Angola  and  east  to 
Uganda.  The  larvae  are  predators  of  Jassidae  and  Membracidae  (Cottrell,  1984).  Gilbert  (1976)  records 
and  illustrates  an  adult  feeding  from  the  'nectary'  of  a  lycaenid  larva. 

Species  limits  within  Megalopalpus  are  very  imperfectly  understood.  Aurivillius  (1922),  followed  by 
Stempffer  (1967)  and  Cottrell  (1984),  accepted  four  species:  metaleucus,  simplex,  zymna  and  angulosus. 
Of  these,  Stempffer  (1967)  dissected  specimens  of  the  first  three,  and  stated  that  'The  male  genitalia  of  M. 
zymna  and  M.  metaleucus  are  very  similar  to  those  of  simplex'.  D'Abrera  (1980)  and  Carcasson  (1981), 
however,  following  Peters  (1952),  recognised  only  three  species,  zymna  (to  include  simplex),  metaleucus 
and  angulosus  -  the  last,  representing  the  species  not  dissected  by  Stempffer,  they  regarded  as  doubtful, 
and  possibly  only  a  synonym  of  zymna.  Berger  (1981)  also  recognised  only  three  species  but,  in  contrast, 
these  were  zymna,  simplex  and  metaleucus  -  no  mention  was  made  of  angulosus  (confounded  by  Berger 
with  metaleucus  -  see  below). 

In  the  BMNH  as  apparently  curated  by  Ms  S.  J.  May,  all  four  species  listed  by  Aurivillius  and  Stempffer 
are  purportedly  represented.  However,  it  is  very  difficult  to  see  how  simplex  differs  consistently  from 
zymna,  and  under  the  former  the  following  note  appears:  'Probably  some  simplex  in  zymna  series,  as  these 
were  hitherto  considered  to  be  synonymous.  SJM.'.  This  note  is  doubly  puzzling  because  not  only  have  the 
two  often  been  considered  to  be  separate,  there  is  no  evidence  of  comprehensive  dissection  of  the 
specimens  segregated  as  simplex  (and  therefore,  presumably,  their  distinguishing  features  must  have  been 
open  to  inspection).  The  basis  of  this  separation,  if  indeed  it  is  valid,  is  thus  unclear  or  unknown.  Carcasson 
(1981)  implies,  effectively,  that  there  are  just  two  species:  large  (metaleucus)  and  small  (zymna). 
Unfortunately,  the  type  of  zymna  is  of  intermediate  size!  However,  there  seems  little  doubt  that  there  are 
at  least  three  species,  corresponding  to  the  arrangement  in  Peters  (1952)  and  illustrated  by  D'Abrera 
(1980).  All  four  nominal  species  are  grossly  sympatric  in  some  areas  (e.g.  Cameroun,  Zaire),  although 
Clench  (1965)  noted  a  'strong  correlation  with  geography'  for  the  four  forms  (including  ?  simplex)  that  he 
described  under  zymna.  Detailed  field  observations  and  reared  material  are  probably  essential  for  the 
solution  of  this  problem. 

In  the  following  key,  simplex  and  zymna  are  separated  following  Aurivillius  (1922)  and  Berger  (1981), 
but  this  is  not  very  reliable  if  the  identifications  in  the  BMNH  are  correct;  see  also  Clench  (1965). 

Key  to  the  species  of  the  genus  Megalopalpus 

1  Larger  species,  forewing  length  17-23  mm;  black  border  of  forewing  upperside  does  not  extend 

broadly  quite  as  far  posterior  as  vein  A\\  catenulate  markings  of  hindwing  underside  darker 

than  general  ground  colour,  not  solely  forming  a  series  of  concentric  bands 2 

Smaller  species,  forewing  length  11-18-5  mm;  black  border  of  forewing  upperside  extends 
broadly  and  fully  to  vein  A\;  catenulate  markings  of  hindwing  underside  not  appreciably 
darker  than  ground  colour,  outlined  by  pale  scales,  forming  a  series  of  roughly  concentric 
bands 3 

2  Hindwing  upperside  very  narrowly  bordered  (up  to  1  mm)  with  dark  scales  along  posterior 

margin;  postdiscal  catenulate  band  of  hindwing  underside  disrupted  medially;  hindwing 

underside  pattern  with  overall  appearance  of  veined  marble metaleucus  (p.  85) 

-    Hindwing  upperside  usually  more  broadly  bordered  (2-3  mm)  with  dark  scales,  affecting  entire 


THE  MILETINI  85 

outer  margin;  postdiscal  catenulate  band  of  hindwing  underside  reticulate  medially;  hindwing 

underside  pattern  with  a  more  complex  marbling  effect angulosus  (p.  85) 

3    Hindwing  upperside  with  dark  marginal  band  3-5  mm  in  width,  not  interrupted  in  middle 

zymna  (p.  85) 

Hindwing  upperside  with  dark  marginal  band  no  more  than  3  mm  in  width  (often  much  less), 
medially  very  narrow  or  completely  interrupted simplex  (p.  85) 

Megalopalpus  angulosus  Griinberg 

Megalopalpus  angulosus  Grunberg,  1910:  478;  D'Abrera,  1980:  469  (illustr.).  3  cf,  3  $  syntypes, 

EQUATORIAL  GUINEA:  Makomo  and  Alcu  (depository  unknown). 
[Megalopalpus  metaleucus  Karsch;  Berger,  1981:  pi.  197,  fig.  5.  Misidentification.] 

This  butterfly,  judging  by  the  underside  and  if  correctly  identified  in  the  BMNH,  is  a  good  species  and  not 
merely  a  maculated  or  seasonal  form  of  metaleucus .  M.  angulosus  is  known  from  Cameroun,  Equatorial 
Guinea  and  Zaire.  Specimens  in  the  BMNH  have  a  forewing  length  of  20-23  mm. 

Megalopalpus  metaleucus  Karsch 

[Allotinus  zymna  (Westwood);  Grose-Smith  &  Kirby,  1891:  figs  1,  2.  Misidentification.] 
Megalopalpus  metaleucus  Karsch,  1893:  217;  Clench,  1965:  326,  figs  198-201;  D'Abrera,  1980:  470 
(illustr.);  Berger,  1981:  pi.  197,  figs  3, 4.  2  cf  syntypes,  TOGO:  Bismarckburg  (7MNHU)  [not  examined]. 

M.  metaleucus  is  considered  to  occur  from  Liberia  and  Ivory  Coast  south  to  Cameroun  and  Zaire,  and  east 
to  Uganda.  Specimens  in  the  BMNH  have  a  forewing  length  range  of  17-22  mm.  Clench  (1965)  discusses  a 
variation  of  the  species. 

Megalopalpus  simplex  Rober 

Megalopalpus  simplex  Rober,  1886:  51,  pi.  4,  fig.  4.  $  syntype(s),  EQUATORIAL  AFRICA:  'Borneo'  (patria 

falsa)  (?SMT)  [not  examined]. 
Liptena  bicoloria  Capronnier,  1889:  121.  Holotype  (sex?),  ZAIRE:  Lopori  Shoven  (Martini)  (depository 

unknown).  [Synonymy  from  Stempffer,  1967.] 
Allotinus  similis  Kirby,  1890:  262.  Syntype(s),  (s'ex?),  CAMEROUN:  Barombi  (Preuss.)  (MNHU)  [not 

examined].  [Synonymy  from  Aurivillius,  1922;  Stempffer,  1967.] 
Megalopalpus gigas  Bethune-Baker,  1914:  335,  pi.  58,  figs  9, 9a.  cf  syntypes,  CAMEROUN/GABON  (BMNH) 

[examined].  [Synonymy  from  Stempffer,  1967.] 

Material  in  BMNH  identified  as  simplex  is  very  variable  with  respect  to  the  hindwing  dark  border  (see  also 
zymna  below).  As  already  discussed,  the  basis  of  the  separation  of  simplex  from  zymna  is  elusive,  and  may 
not  be  real.  The  identification  of  specimens  illustrated  by  Berger  (1981)  to  represent  both  species  should  be 
regarded  with  caution.  The  forewing  length  of  the  BMNH  material  varies  from  14-0-18-5  mm.  The 
distributional  range  is  thought  to  include  Ghana,  Liberia,  Nigeria,  Cameroun,  Gabon,  Zaire  and  Uganda. 

Megalopalpus  zymna  (Westwood) 

Pentila  zymna  Westwood,  1851:  pi.  76,  fig.  7;  1852:  503.  1  cf  syntype,  NIGERIA:  Ashanti  (ex  Wesleyan 

Missionary  Society)  (BMNH)  [examined]. 
Megalopalpus  zymna  f.  pallida  Aurivillius,  1922:  362.  Holotype  cf ,  UGANDA:  Ruwenzori  (?Stockholm) 

[not  examined]. 

As  already  noted,  it  is  doubtful  if  simplex  can  be  reliably  separated  from  zymna.  However,  Clench  (1965) 
recognised  four  more  or  less  distinct  forms  under  zymna,  including  one  apparently  corresponding  to 
simplex,  and  the  possibility  that  zymna  s.l.  represents  a  species  complex  must  be  considered.  Specimens 
identified  in  the  BMNH  as  zymna  have  a  forewing  length  of  11-18  mm,  with  the  lone  syntype  at  18-5  mm 
(despite  this,  the  syntype  is  definitely  of  the  correct  general  facies  -  it  is  not  angulosus  or  metaleucus  as 
currently  understood).  The  distribution  of  zymna  encompasses  that  given  for  simplex,  and  is  considered  to 
extend  also  to  Macias  Nguema  (Fernando  Poo),  Equatorial  Guinea,  Angola,  southern  Sudan  and 
Zimbabwe. 


86  J.  N.  ELIOT 

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Staudinger  and  Mr  Henley  Grose  Smith.  Annals  and  Magazine  of  Natural  History  (6)  6:  261-274. 

Lewis,  H.  L.  1974  Butterflies  of  the  World  xvi  +  104  pp.,  208  pis.  London. 

Moore,  F.  1857.  Catalogue  of  the  lepidopterous  insects  in  the  Museum  of  the  Honourable  East  India 
Company  5  +  278  +  4  +  11  pp.,  12  +  6 pis.  London. 

—  1866.  On  the  lepidopterous  insects  of  Bengal.  Proceedings  of  the  Zoological  Society  of  London  1865: 
755-823,  3  pis. 

-  1883.  Descriptions  of  new  Asiatic  diurnal  Lepidoptera.  Proceedings  of  the  Zoological  Society  of 
London  1883:  521-535,  2  pis. 

-  1884.  Descriptions  of  some  new  Asiatic  diurnal  Lepidoptera;  chiefly  from  specimens  in  the  Indian 
Museum,  Calcutta.  Journal  of  the  Asiatic  Society  of  Bengal  (2)  53:  16-52. 

1886.  List  of  the  Lepidoptera  of  Mergui  and  its  Archipelago  collected  for  the  Trustees  of  the  Indian 


Museum,  Calcutta,  by  Dr  John  Anderson  F.R.S.,  Superintendent  of  the  Museum.  Journal  of  the 
Linnean  Society  of  London  (Zoology)  21:  29-60,  2  pis. 

Moulton,  J.  C.  1911.  A  list  of  the  butterflies  of  Borneo  with  descriptions  of  new  species  (3).  Journal  of  the 
Straits  Branch  of  the  Royal  Asiatic  Society  60:  73-177, 1  pi. 

Murayama,  S.  &  Okamura,  H.  1973.  Butterflies  of  Luzon  Island  in  the  Philippines,  with  descriptions  of 
new  species  and  subspecies.  Tyo  to  Ga  24:  10-25, 48  figs. 

Okubo,  K.  1983.  Butterflies  of  Tioman  Island,  West  Malaysia,  with  the  description  of  new  species.  Tyo  to 
Ga  33:  168-184, 47  figs. 

Opinion  566  1959.  Suppression  under  the  Plenary  Powers  of  the  generic  name  Tingra  Boisduval,  1847,  and 
designation  under  the  same  Powers  for  the  genera  Liptena  Westwood,  [1851],  and  Pentila  Westwood, 
[1851],  of  type  species  in  harmony  with  accustomed  usage  (Class  Insecta,  Order  Lepidoptera).  Opinions 
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377-390. 

Peters,  W.  1952.  A  provisional  check-list  of  the  butterflies  of  the  Ethiopian  region  201  pp.  Feltham,  Middx. 

Piepers,  M.  C.  &  Snellen,  P.  C.  T.  1918.  The  Rhopalocera  of  Java  Erycinidae,  Lycaenidae.  xiv  +  114  pp. ,  9 
pis.  The  Hague. 

Pinratana,  A.  1981.  Butterflies  in  Thailand  4:  vii  +  215  pp.,  36  pis,  47  figs.  Bangkok. 

Reuter,  E.  1897.  Uber  die  Palpen  der  Rhopaloceren.  Acta  Societatis  Scientiarum  Fennicae  22:  xi  +  588  pp. , 
6  pis. 

Ribbe,  C.  1889.  Beitrage  zur  Lepidopteren-Fauna  von  Gross-Ceram.  Deutsche  Entomologische  Zeit- 
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-  1926.  Neue  Lycaenenformen  hauptsachlich  von  Celebes.  Entomologische  Mitteilungen  15:  78-91. 
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Royal  Entomological  Society  of  London  94:  247-271,  2  pis. 

Riley,  N.  D.  &  Godfrey,  E.  J.  1921.  Some  undescribed  Rhopalocera  from  Siam.  Journal  of  the  Natural 
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Rober,  J.  1886.  Neue  Tagschmetterlinge  der  Indo-Australischen  Fauna.  Deutsche  Entomologische  Zeit- 
schrift, Iris  1:  45-72,  4  pis. 

-  1892.  In  Staudinger,  O.  &  Schatz,  E.,  Exotische  Schmetterlinge  2.  Die  Familien  und  Gattungen  der 
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Roepke,  W.  1918.  Zur  Myrmekophilie  von  Gerydus  boisduvali  Moore.  Tijdschrift  voor  Entomologie  61: 

1-16,  3  pis,  2  figs. 
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1932.  Over  eenige  nieuwe  of  weinig  bekende  vlindersoorten.  Verslagen  van  de  Vergaderingen  der 

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251-534,  pis  31-80  +  suppl.  pi.  London. 


90  J.  N.  ELIOT 


Fig.  55  Allotinus  (Allotinus)  agnolia  sp.  n.,  $  paratype;  Sumatra. 

Fig.  56  Allotinus  (Allotinus)  nicholsi  battakanus  Fruhstorfer,  cf  lectotype;  Sumatra. 

Fig.  57  Allotinus  (Allotinus)  nicholsi  battakanus  Fruhstorfer,  $  paralectotype;  Sumatra. 

Fig.  58  Allotinus  (Fabitaras)  borneensis  Moulton,  cf ;  Borneo. 

Fig.  59  Allotinus  (Fabitaras)  punctatus  $-f.  caesemius  Fruhstorfer;  Mindanao. 

Fig.  60  Allotinus  (Fabitaras)  nigritus  Semper,  $ ;  Mindanao. 

Fig.  61  Allotinus  (Fabitaras)  strigatus  strigatus  Moulton,  d" ;  Borneo. 


THE  MILETINI 


91 


61 


92  J.  N.  ELIOT 


Fig.  62  Allotinus  (Fabitaras)  bidiensis  sp.  n. ,  C?  holotype;  Borneo. 

Fig.  63  Allotinus  (Fabitaras)  brooksi  sp.  n. ,  cf  holotype;  Borneo. 

Fig.  64  Allotinus  (Paragerydus)  melos  (H.  H.  Druce),  cf  topotype;  Cagayan  I. 

Fig.  65  Allotinus  (Paragerydus)  samarensis  samarensis  sp.  n. ,  $ ;  Mindanao. 

Fig.  66  Allotinus  (Paragerydus)  samarensis  russelli  subsp.  n.,  $  holotype;  Sulawesi. 

Fig.  67  Allotinus  (Paragerydus)  macassarensis  macassarensis  (Holland),  cf ;  Sulawesi. 

Fig.  68  Allotinus  (Paragerydus)  macassarensis  macassarensis  (Holland),  $ ;  Sulawesi. 


THE  MILETINI 


93 


64 


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94  J.   N.  ELIOT 


Fig.  69  Allotinus  (Paragerydus)  luzonensis  Eliot,  cf  holotype;  Luzon. 

Fig.  70  Allotinus  (Paragerydus)  luzonensis  Eliot,  9  allotype;  Luzon. 

Fig.  71  Allotinus  (Paragerydus)  albatus  albatus  C.  &  R.  Felder,  $  holotype;  Sulawesi. 

Fig.  72  Allotinus  (Paragerydus)  albatus  mendax  subsp.  n.,  cf  paratype;  Luzon. 

Fig.  73  Allotinus  (Paragerydus)  albatus  mendax  subsp.  n.,  9  allotype;  Luzon. 

Fig.  74  Allotinus  (Paragerydus)  apries  apries  Fruhstorfer,  cf ;  Borneo. 

Fig.  75  Allotinus  (Paragerydus)  nivalis  felderi  Semper,  cf ;  Mindanao. 

Fig.  76  Allotinus  (Paragerydus)  substrigosus  ballantinei  subsp.  n. ,  cf  holotype;  Palawan. 


THE  MILETINI 


95 


70 


74 


75 


96  J.  N.  ELIOT 


Fig.  77  Allotinus  (Paragerydus)  substrigosus  yusukei  subsp.  n. ,  cf  holotype;  Mindanao. 

Fig.  78  Logania  nehalemla  Fruhstorfer,  cf  holotype;  New  Guinea. 

Fig.  79  Logania paluana  sp.  n.,  $  holotype;  Sulawesi. 

Fig.  80  Logania  obscura  Rober  subsp. ,  cf ;  Banggai  I. 

Fig.  81  Logania  distanti  distanti  Semper,  cf ;  Luzon. 

Fig.  82  Lontalius  eltus  eltus  sp.  n. ,  $  holotype;  Pulo  Laut. 

Fig.  83  Miletus  mallus  mallus  (Fruhstorfer),  cf ;  Vietnam:  south. 

Fig.  84  Miletus  gaesa  gaesa  (de  Niceville) ,  cf  var. ;  West  Malaysia. 

Fig.  85  Miletus  nymphis  eneus  Eliot,  cf ;  Sumatra:  south-west. 


THE  MILETINI 


97 


78 


82 


85 


98  J.  N.   ELIOT 


Fig.  86  Miletus  cellarius  (Fruhstorfer),  cf ;  Borneo:  Kina  Balu. 

Fig.  87  Miletus  cellarius  (Fruhstorfer),  $ ;  Borneo:  Kina  Balu. 

Fig.  88  Miletus  symethus  near  subsp.  phantus  subsp.  n. ,  9  5  Luzon. 

Fig.  89  Miletus  symethus  hierophantes  (Fruhstorfer),  cf ;  Sulu  Is. 

Fig.  90  Miletus  symethus  near  subsp.  hierophantes  (Fruhstorfer),  cf ;  Mindanao. 

Fig.  91  Miletus  symethus  near  subsp.  hierophantes  (Fruhstorfer),  $ ;  Mindanao. 

Fig.  92  Miletus  atimonicus  Murayama  &  Okamura,  cf ;  Negros. 

Fig.  93  Miletus  atimonicus  Murayama  &  Okamura,  9 ;  Luzon. 


THE  MILETINI 


99 


100 


J.  N.  ELIOT 


94 


Figs.  94-98  94,  Miletus  heradeion  near  subsp.  heradeion  (Doherty),  cf ;  Sarawak;  95  Miletus  heradeion 
near  subsp.  heradeion  (Doherty),  $ ;  Borneo:  Pulo  Laut;  96,  Miletus  heradeion  arion  Eliot,  cf ;  Borneo: 
Kina  Balu;  97,  Miletus  melanion  melanion  cf-f.  melanion  C.  &  R.  Felder,  cf;  Leyte;  98,  Miletus 
melanion  melanion  cf-f.  albiguttatus  f.  n.,  cf  holotype;  Luzon. 


THE  MILETINI 


101 


99 


102 


100 


103 


104 


Figs.  99-104  99,  Miletus  bazilanus  (Fruhstorfer) ,  O";  Mindanao;  100,  Miletus  bazilanus  (Fruhstorfer),  $ 
paralectotype;  Bazilan;  101,  Miletus  bazilanus  (Fruhstorfer),  $  var.;  Mindanao;  102,  Miletus  takanamii 
sp.  n. ,  cf  holotype;  Mindanao;  103,  Miletus  takanamii  sp.  n. ,  $  allotype;  Mindanao;  104,  Miletus  celinus 
Eliot,  cf ;  Sulawesi. 


102 


J.  N.  ELIOT 


106 


Fig.  105    Allotinus  (Paragerydus)  samarensis  samarensis  sp.  n. ,  cf  holotype;  Samar. 
Fig.  106    Allotinus  (Paragerydus)  apries  ristus  subsp.  n.,  d"  holotype;  Palawan. 


THE  MILETINI 


103 


107 


Fig.  107     Logania  waltraudae  sp.  n.,  C?  holotype;  Samar. 

Fig.  108     Lontalius  eltus  treadawayi  subsp.  n.,  $  holotype;  Samar. 


104 


J.  N.  ELIOT 

Index 


Principal  references  are  in  bold;  synonyms,  infrasubspecific  and  unavailable  names  are  in  italics 


absens  30 
acampsis  5 
acragas  5 
acrisius  5 
adeus5 

agnolia  3,  8,  15 
albatus3,32,41 
albifasciatus3,  8,  13 
albiguttatus  5  ,  80 
albotignula  5 
alkamah  3,  14 
Allotinus3,  7 
amphiarus  5 
anaxandridas  3,  23 
ancius3,  11 
ancon5,  77 
anconides  5 
angulosus  5,  85 
aphacus3,  10,  11 
aphocha3,48 
aphthonius  4,  56,  57 
aphytis  5 

apries3,31,42,  43 
apsines  4,  70 
apus3,  10,  12 
Archaeogerydus  4,  75 
archilochusS,  77 
arionS,  80 
aronicus  5 
arrius3,  21 
artaxatus  4 
artinus  3,  11 
assamensis  4 


atimonicus  5,  77,  79 
atomaria  5 
audax3,  10,  12 
avitus  5 

bajanus  4,  49 
ballantinei  4,  53,  55 
bangkanus  5 
battakanus3,  16,  17 
batuensis,  Allotinus  3,  35,  36 
batuensis,  Miletus  5 
batunensis  5 
bazilanus5,77,82 
bicoloria5,  85 
bidiensis3,20,  26 
biggsiiS,  77 
boisduvali  Moore  5,  77 
boisduvalii  Butler  5 
borneensis3,20,  22 
brooksi3,20,  26 
buruensis  5 


celinus  5,  76,  77 
ceramensis  5 
chinensis  4,  76 
cineraria  4,  65 
continentalis  3,  46,  47 
corbeti3,31,44 
corus  4 
courvoisieri  5 
croton  4,  76 

damis  4,  64,  65 

damodar3,  40 

davidis  4,  32,  55 

denalus3,25 

denticulata  4 

depictus  3,  18 

diehli  4,  64,  66 

dilutus3,47 

diopeithes  5 

diotrophes  5 

distant!  Semper  4,  59,  69,  70 

distanti  Staudinger  4,  67 

divisa  5 

donussa  4,  68 

dositheus3,29,42,44 

dossemus  5 

dotion  3,  10,  12 

drucei,  Logania  4,  70,  71 

drucei,  Miletus  5,  77 

drumila4,  31,  56 

edonus  5 
elioti3,22 

eltus  4,  74 
enatheus  4,  49 
eneus  4,  76 
enganicus  4,  49 
epldurus  5 
eretria  3,  24 
eryximachus  3,  10,  11 


,23 
carrinas  4 
catoleucos  5 
caudatus  3,  21 


eupalion  3,  43 
euphranor5,81,82 
eurytanus  3,  47 
eustatius  4 
evora  4,  63 
extraneus  5 

Fabitaras3,7,  20 
fabius3,20,21 
fallax3,8,  10 
faustina4,  65,68 
felderi  4,  52 
fictus  4 
florensis  5 
fruhstorferi3,  30 

gaesa  4,  76 
gaetulusS,  77 
gallus5,77 
gardineri  5 
georgius  4,  46,  49 


Gerydinae  1 

Gerydus  4,  75 

gethusus4,  78 

gigantes5;77,79 

g/gos  Bethune-Baker  5,  85 

gigas  H.  H.  Druce  5 

glypha  4,  67 

gopara  4, 77 

grisea  4,  57 

hampsoni  4,  59,  72 
heracleion5,76,  77,80 
heraeon  5 
hilaeira4,  64,  65 
hierophantes  5,  78 
hieropous  5 
horsfieldi3,31,32,33 
hyllus  5 

improbus  4 
infumata3,34 
innocens  5 
ins  ignis  4, 57 
intricata3, 35 
irroratus  4 

jacchus  5 
javanica  4,  64, 67 

kalawarus  3, 18 
kallikrates  3, 14 
karennius  4 
kelantanus  4 

Iahomius4,  65,  66 
learchus  4 
/e/fws  4, 49 
Ienaia4,  53,  54 
leogoron  3,  32,  34,  35 
Ieos5,77 
leucocyon  5 
Iindus3,34,35 
Logania  4,  7,  57 
lombokianus  5 
longeana  4 
Lontalius  4,  7,  74 
luca  4, 71 
Iuzonensis3,  32,40 

macassarensis  3,  31, 39 
magaris  4,  53 
maitus3,29 
major  3,  8, 17 
Malais  4,  57 
malayanus3,  25 
malayica  4,  59,  60 
mallus  4,  76,  78 
mangolicus  5 
manychus  3, 14 
marmorata  4,  59,  64,  65 
martinus  4,  68 


masana  4,  72 
massalia  4,  70 
maximus,  Allotinus  3,  8,  19 
maximus,  Miletus  5 
meeki  4,  72 
Megalopalpus  5,  7,  84 
megaris  5 

melanion5,77,80,  81 
melos3,  32,  36 
menadensis  3,  39,  40 
mendava  3,  28 
mendax  3,  41 
meronus  5 
metaleucus  5,  84,  85 
metrovius  5 
michaelis  3,  12 
Miletographa  3,  30 
Miletus  4,  7,  75 
milvius  4 
minis  3,  14,  15 
molionides  3  ,  48 
moorei  3,  46,  48 
multistrigatus  4,  57 
munichya  4,  64,  66 
myriandus  4,  48 


,  65 

narsares  3,  29 
natunensis  5 
nehalemia  4,  59,  60 
ness  us  3,  34 
niasicus  5 
nicer  atus  4,  49 
nicholsi3,8,  16,  17 
nigritus3,20,  24 
nineyanus  5 
nivalis  4,  32,51,52 
normani3,  35,36 
nuctus  5 
nymphis  4,  76 

obscura  Distant  &  Pryer  4,  65 
obscuraR6ber4,59,68 
oichalia  5 
oxylus  5 

paetus  4,  32,  50 


INDEX 

paianlus  5 
pallaxopas  5 
pallida5,8S 
paluana  4,  59,  63 
palawana  4,  64, 67 
pamisus  3,  21 
pandu 5 
panormis  3,  27 
Paragerydus  3,  7,  30 
parapus4,  31,  51 
pardus  4 
pentheus  5 
perlucidus  5 
permagnus  3, 33 
petronius  5 
phantusS,  78 
philippus  5 
philopator  5 
phradimon  5 
plessis3,  35, 36 
porriginosus3,  28 
portunus3,  21,  29 
porus  4 
posidion  3,  48 
punctatus3,20,  23 
pyxus3,29,30 

rebilus  3, 48 
regina  4,  59,  62, 63 
rekkia  3, 46,  48 
reverdini3,  36 
ristus3,42,44 
russelli3,  38 

sabazus3, 10 
samarensis  3,  31,  37,  38 
samosata4,  65,67 
sarrastes3,  21,  28 
sarus  5 

satelliticus3,  33,  34 
sebethus  5 
shanius  4 
siamensisS,  79 
sibyllinus4,  53,  54 
silarus  3, 14 
simalurensis  5 
similis  5,  85 


105 


simplex  5,  85 
siporanusS,  33,34 
solitarius5,78 
sora  4,  65 
sriwa  4,  62,  63 
staudingeri  4,  70,  72 
stenosa  4,  65 
strigatus3,20,  24,  25 
stygianus  5 
subfasdata  4,  73 
substrigosus  4,  32,  53 
subura  4,  60 
subviolaceus  3,  8, 14 
suka  4,  49 
Symetha  4,  75 
symethusS,  77,  78 

takanamii  5,  77,  83 

talu3,36 

taras3,21,27 

tavoyana  4 

tellus  5 

teos5 

treadawayi  4,  74 

turdeta  4, 70 

tymphrestus  3, 10, 12 

unicolor  3,  32,  45,  46,  47 

vadosus3,35 
valeus5,77 
vaneeckei  5 
vespesianus  5 
vincula  5 
virtus  5 
vitelianus  5 , 82 

waltraudae  4,  59,  61 
waterstradti  3,  30 
watsoniana  4,  59,  73 

xeragis  5 
yusukei4,  53,  55 

zaradrus3,  10 
zinckenii  4,  77 
zitema  4,  46,  50 
zymnaS,  85 


Occasional  Papers  on  Systematic  Entomology 

New  Series 


The  economic  importance  of  insects,  and  the  enormous  number  of  species,  have  resulted  in  a 
vast  literature  written  in  many  languages;  that  which  is  of  direct  economic  importance  and 
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No.  1.  A  checklist  of  Neotropical  arctiine  and  pericopine  tiger  moths. 

A.  Watson  &  D.  T.  Goodger 

72pp.  inc.  4  colour  plates    27  February  1986 

No.  2.  An  annotated  checklist  of  the  Carabidae  (including  Cicindelinae,  Rhysodinae  and 

Paussinae)  recorded  from  Borneo. 

N.  E.  Stork 

26pp. ,  1  map    29  May  1986 


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A  review  of  the  Miletini  (Lepidoptera:  Lycaenidae) 

By  J.N.Eliot 

Australian  ichneumonids  of  the  tribes  Labenini  and  Poecilocryptini 

By  I.  D.  Gauld  &  G.  A.  Holloway 

The  tribe  Pseudophloeini  (Hemiptera:  Coreidae)  in  the  Old  World  tropics  with  a  discussion  on 
the  distribution  of  the  Pseudophloeinae 

By  W.  R.  Dolling 

The  songs  of  the  western  European  grasshoppers  of  the  genus  Omocestusin  relation  to  their 
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Australian  ichneumonids  of  the  tribes 
Labenini  and  Poecilocryptini 

I.  D.  Gauld  &  G.  A.  Hollo  way 


Entomology  series 

Vol53  No  2  30  October  1986 


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Entomology  series 
Vol  53  No  2  pp  107-150 


Issued  30  October  1986 


D  r?  *"  • 

Australian  ichneumonids  of  the  tribes  Labenini  and 


Poecilocryptini 


I.  D.  Gauld 

Department  of  Entomology,  British  Museum  (Natural  History),  Cromwell  Road,  London 
SW7  5BD 

G.  A.  Holloway 

Division  of  Invertebrate  Zoology,  Australian  Museum,  6-8  College  Street,  Sydney,  New 
South  Wales,  Australia 

Contents 

Synopsis 107 

Introduction : 107 

Material  examined 108 

Checklist  of  Australian  Labenini  and  Poecilocryptini 108 

Subfamily  Labeninae 109 

Key  to  tribes  of  Labeninae 110 

Tribe  Labenini 110 

Key  to  genera  of  Labenini  occurring  in  Australia Ill 

Labena  Cresson Ill 

Key  to  species  of  Labena  occurring  in  Australia 112 

Certonotus  Kriechbaumer 117 

Key  to  species  of  Certonotus  occurring  in  Australia 119 

Tribe  Poecilocryptini 137 

Key  to  genera  of  Poecilocryptini 138 

Alaothrys  Gauld 138 

Poecilocryptus  Cameron 139 

Key  to  species  of  Poecilocryptus 139 

Urancyla  Gauld 141 

Acknowledgements 142 

References 142 

Index  to  hosts 149 

Index  to  Ichneumonidae 149 

Synopsis 

The  Australian  species  of  the  labenine  tribes  Labenini  and  Poecilocryptini  are  revised  and  keys  provided  to 
the  five  genera  and  36  species  occurring  on  the  continent.  Twenty  new  species  are  described,  but  one  is  not 
formally  named  as  its  status  requires  further  investigation.  The  remaining  16  species  are  redescribed  and 
their  diagnostic  features  emphasized.  Asperellus  Townes  is  newly  placed  as  a  junior  synonym  of  Certonotus 
Kriechbaumer,  Certonotus  tasmaniensis  Turner  is  treated  as  a  junior  synonym  of  C.  nitidulus  Morley  and 
Poecilocryptus  straminea  Morley  placed  as  a  junior  synonym  of  P.  nigromaculatus  Cameron.  Lectotypes 
are  designated  for  five  species.  Details  of  known  hosts  are  given  together  with  notes  about  the  geographical 
distribution  of  each  species.  A  brief  introductory  section  discusses  the  systematic  position  of  the  subfamily 
and  outlines  possible  relationships  between  species.  A  checklist  of  Australian  species,  and  indexes  to  hosts 
and  parasitoids  complete  the  work. 

Introduction 

Amongst  the  most  important  of  the  natural  enemies  of  insect  pests  are  the  parasitic  Hymenop- 
tera,  a  very  large  group  of  animals  whose  larvae  develop  at  the  expense  of  other  insects  (Askew, 
1971).  Under  normal  circumstances,  the  populations  of  many  injurious  insects  are  severely 


Bull.  Br.  Mus.  nat.  Hist.  (Ent.)  53  (2):  107-149  Issued  30  October  1986 


108  I.  D.  GAULD  &  G.  A.  HOLLOWAY 

limited  by  the  attacks  of  Parasitica,  and  in  several  countries,  including  Australia,  the  ravages  of 
accidentally  imported  pests  have  been  curtailed  by  the  introduction  of  one  or  more  hymenopter- 
ans  (Muldrew,  1967;  Taylor,  1978).  Other  introduced  pests  have  been  severely  attacked  by 
native  Australian  parasitoids  (Tryon,  1900).  During  the  past  30  years  there  has  been  an  upsurge 
in  interest  in  using  Hymenoptera  and  other  organisms  for  purposes  of  pest  control  (Wilson, 
1960;  Huffaker  &  Messenger,  1976)  as  an  alternative  to  costly,  ineffective  and  environmentally 
destructive  chemical  methods  (Bosch,  1978).  However,  for  biological  control  programmes  to  be 
successful  an  intimate  knowledge  is  necessary  of  the  life  history  and  interactions  of  the  pest  and 
its  parasites.  A  sound  taxonomic  basis  is  vital  for  the  development  of  such  knowledge  (Hardy, 
1982),  for  such  work  permits  the  accurate  identification  of  an  organism  and  hence  provides 
constancy  and  universality  in  the  usage  of  names,  a  prerequisite  for  the  national  and  internation- 
al communication  of  information. 

The  present  work  is  a  taxonomic  study  of  two  very  distinct  tribes  of  one  of  the  apparently  most 
primitive  extant  ichneumonid  subfamilies,  the  Labeninae.  These  two  tribes,  the  Labenini  and 
Poecilocryptini,  are  virtually  confined  to  the  southern  hemisphere  with  most  species  occurring  in 
Australia  and  South  America.  The  generic  phylogeny  of  the  Labeninae  and  their  geographical 
distribution  has  recently  been  studied  (Gauld,  1983)  but  no  keys  are  currently  available  to 
facilitate  identification  of  the  species.  This  paper  is  an  attempt  to  provide  a  means  of  determining 
the  Australian  species  of  this  interesting  group. 

The  terminology  in  this  work  follows  that  of  Gauld  (1984). 

Material  examined 

The  study  is  based  on  examination  of  almost  all  specimens  available  in  collections  of  Australian 
ichneumonids.  Special  attention  was  paid  to  collections  in  agricultural  institutions  that  contain  a 
large  number  of  reared  specimens.  Examination  of  these  collections  has  been  supplemented  by 
extensive  collecting,  particularly  in  Tasmania  and  the  south-east.  Although  the  resulting  sample 
is  thought  to  be  fairly  representative  of  the  fauna  of  the  more  humid  eastern  part  of  the 
continent,  relatively  little  material  has  been  examined  from  the  west. 

The  following  abbreviations  have  been  used  for  museums  containing  Australian  material. 
AM  Australian  Museum,  Sydney,  New  South  Wales,  Australia 

ANIC  Australian  National  Insect  Collection,  Canberra,  Australian  Capital  Territory,  Australia 

BMNH  British  Museum  (Natural  History),  London,  U.K. 

BPBM  Bernice  P.  Bishop  Museum,  Honolulu,  Hawaii,  U.S.A. 

MNHN  Museum  National  d'Histoire  Naturelle,  Paris,  France 

MNHU  Museum  fur  Naturkunde  der  Humboldt-Universitat,  Berlin,  D.D.R. 

NMV  National  Museum  of  Victoria,  Melbourne,  Victoria,  Australia 

NSWDA  New  South  Wales  Department  of  Agriculture,  Rydalmere,  New  South  Wales,  Australia 

QM  Queensland  Museum,  Brisbane,  Queensland,  Australia 

TC  Townes  Collection ,  Ann  Arbor ,  Michigan ,  U .  S .  A . 

WAD  A  Western  Australian  Department  of  Agriculture,  Perth,  Western  Australia,  Australia 


Checklist  of  Australian  Labenini  and  Poecilocryptini 


LABENINI 

CERTONOTUS  Kriechbaumer 
Asperellus  Townes  syn.  n. 
andrt-wisp.  n. 
annulatus  Morley 
apicalis  Morley 
iivi/u.vsp.  n. 
ce/eussp.  n. 
cestussp.  n. 
farrugiaisp.  n. 
geniculatus  Morley 
hinnuleus  Krieger  comb.  rev. 


humeralifer  Krieger 

ixion  sp.  n. 

leeuwinensis  Turner  comb.  rev. 

mogimbensis  Cheesman  comb.  rev. 

monticola  Morley 

nitidulus  Morley 

tosmaniensis  Turner  syn.  n. 
palunia  sp.  n. 
pineussp.  n. 
rufescens  Morley 
sisyphus  sp.  n. 
talus  sp.  n. 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  109 

toolangisp.  n.  POECILOCRYPTINI 

zebrus  sp.  n.  ALAOTHYRIS  Gauld 

species  A  elongissimus  Gauld 

LABENA  Cresson  POECILOCR  YPTUS  Cameron 

annulata  (Brulle)  coloratussp.  n. 

chadwickii  (Parrott)  galliphagus  sp.  n. 

grand  is  sp.  n.  nigripectus  Turner  &  Waterston 

jacunda  sp.  n.  nigromaculatus  Cameron 

keira  sp.  n.  straminea  Morley  syn.  n. 

malecasta  sp.  n.  URANCYLA  Gauld 

pudenda  sp.  n.  fulva  Gauld 

Nomen  dubium 

Certonotus  varius  Kriechbaumer 

From  Kriechbaumer's  original  description  (1889:  308)  it  is  apparent  that  this  species  is  one  of  the  larger 
Australian  Certonotus.  As  we  have  seen  no  species  that  agrees  with  the  description,  nor  can  the  type  be 
located,  we  have  no  option  but  to  treat  this  as  an  unrecognized  taxon. 

Subfamily  LABENINAE 

The  Labeninae  is  a  moderately  large  subfamily.  Amongst  the  ichneumonid  subfamilies  it  is 
unique  in  being  a  Gondwanic  group.  Most  genera  and  species  are  confined  to  Australasia  and/or 
South  America;  only  a  dozen  or  so  taxa  occur  outside  this  area  (Table  1).  All  other  ichneumonid 
subfamilies  are  well  represented,  if  not  actually  most  diverse,  in  the  Holarctic  region.  Phy- 
logenetic  analysis  suggests  that  the  Labeninae  radiated  in  West  Gondwanaland,  prior  to  the 
separation  of  Australia,  Antarctica  and  South  America,  some  55  mya  (Gauld,  1983).  The  group 
is  only  represented  by  some  highly  derived  species  in  India  and  South  Africa,  suggesting  that  the 
Gondwanic  radiation  occurred  after  the  separation  of  these  continents  (c.  100  mya).  Many 
labenines  are  associated  with  the  southern  temperate  rain  forests,  the  areas  floristically 
dominated  by  Nothofagus,  Araucaria  and  Podocarpus.  One  labenine  genus  is  known  to  oviposit 
into  the  seeds  of  an  Araucaria  in  Australia. 

Structurally  labenines  exhibit  many  of  the  plesiomorphic  features  of  ichneumonids,  and  the 
larvae  are  particularly  primitive  (Short,  1978).  Biologically  they  also  show  many  of  the  so-called 
primitive  features  of  ichneumonids  -  they  are  ectoparasitoids  and  many  are  known  to  parasitize 
wood-boring  insects. 

It  is  not  clear  what  the  sister-group  to  the  labenines  is.  The  more  primitive  taxa  (Labenini) 
closely  resemble  some  Pimplinae,  but  all  the  similarities  can,  we  believe,  be  interpreted  as  either 
symplesiomorphies,  or  parallel  adaptations  to  the  wood-boring  habit.  We  suggest  that  the 
shared  specializations  shown  by  labenines  and  rhyssines  are  parallelisms  rather  than  synapomor- 
phies  because  of  the  striking  differences  in  oviposition  behaviour  between  the  groups.  Labe- 
nines manipulate  the  ovipositor  with  a  specialized  guide  on  the  antero-medial  surface  of  the  hind 
coxa;  rhyssines  achieve  similar  control  with  specialized  claspers  on  the  ventral  surface  of  the 
gaster.  It  is  not  impossible  that  these  taxa  share  a  close  common  ancestor  (a  wood-borer  with  no 
specialization  to  control  the  ovipositor),  but  there  is  no  conclusive  phylogenetic  evidence  to 
support  such  speculation.  We  suggest  that  the  labenines  are  a  primitive  offshoot  of  the  earliest 
ichneumonid  radiation,  and  that  at  an  early  stage  in  their  evolutionary  history  they  became 
isolated  on  a  southern  continent  prior  to  undergoing  a  modest  radiation,  that  has  in  some 
respects  paralleled  that  of  other  ectoparasitic  ichneumonids  (see  Gauld,  1983;  1984). 

The  Labeninae  comprises  four  morphologically  distinctive  tribes,  Labenini,  Groteini,  Poeci- 
locryptini  and  Brachycyrtini,  all  of  which  are  well  represented  in  Australia  (Table  1).  Both  the 
Groteini  and  Brachycyrtini  include  some  genera  for  which  relatively  insufficient  material  is  at 
hand  to  justify  undertaking  a  taxonomic  revision  at  present.  Quite  good  series  of  some  species  of 
Labenini  and  Poecilocryptini  are  available  for  systematic  study,  and  as  there  are  far  more  species 
than  the  few  described  the  opportunity  has  been  taken  to  monograph  these  taxa. 


110  I.  D.  GAULD  &  G.  A.  HOLLO  WAY 

Table  1    The  distribution  of  the  Labeninae. 


1                     8 

1 

5 

S3 

I 

t/5 
O 

—S                             c 

cS 
1 

1 

•5b 

«                    2 

B                  E 

z          S          £ 

1 

< 

1 

co 

1 

z 

1                   1 

u        6 

Xenothyris 

—         —         — 



2 

— 



Labena 

—         —         — 

7 

50 

2 

—        — 

Certonotus 

1              2             9 

23 

5 

— 

—        — 

Apechoneura 

—           —           — 

— 

25 

— 

—        — 

Labium 

1 

50 

1 

— 

—        — 

Grotea 

—           —           — 

— 

20 

3 

—        — 

Macrogrotea 

—           —           — 

— 

10 

— 

—        — 

Alaothyris 

—           —           — 

1 

— 

— 

—        — 

Urancyla 

—           —           — 

1 

— 

— 

—        — 

Poecilocryptus 

—           —           — 

4 

— 

— 

—        — 

Pedunculus 

—           —           — 

— 

5 

— 

—        — 

Adelphion 

2 

6 

— 

— 

—        — 

Monganella 

—           —           — 

1 

?1 

— 

—        — 

Habryllia 

—           —           — 

— 

4 

— 

—        — 

Brachycyrtus 

j 

3 

5 

2 

2             2 

Key  to  tribes  of  Labeninae 

1  Mandible  long  and  slender,  labrum  very  large,  almost  as  long  as  clypeus  (Fig.  1);  female  with 

ovipositor  barely  projecting  beyond  apex  of  gaster ,  dorsoventrally  depressed GROTEINI 

-  Mandible  rather  short  and  not  particularly  slender,  labrum  from  moderately  small  to  virtually 

concealed  (Fig.  2);  female  with  ovipositor  projecting  beyond  apex  of  gaster  by  at  least  0-5 
times  length  of  hind  tibia 2 

2  Fore  wing  with  abscissa  of  Cu\,  between  Im-cu  and  C«la  at  least  1-4  times  as  long  as  Cuib  (Fig. 

3);  hind  coxa  of  female  with  a  furrow  on  anterior  surface  internally;  base  of  valvula  3  bearing 
distinct  sclerotized  lobe  (Fig.  4) LABENINI(p.  110) 

-  Fore  wing  with  abscissa  of  Cui  between  Im-cu  and  Cwla  from  subequal  to,  to  conspicuously 

shorter  than  Cwlb  (Figs  5,  6);  hind  coxa  of  female  without  a  furrow  on  anterior  surface 
internally;  base  of  valvula  3  without  a  distinct  sclerotized  lobe 3 

3  Fore  wing  with  2m-cu  with  one  long  bulla,  sometimes  with  an  indistinct  trace  of  a  vein  centrally 

(Fig.  5);  occipital  carina  dorsally  absent POECILOCRYPTINI  (p.  137) 

-  Fore  wing  with  2m-cu  with  two  short  bullae  widely  separated  from  each  other  (Fig.  6) ;  occipital 

carina  dorsally  complete BRACHYCYRTINI 

The  Groteini  and  Brachycyrtini  are  not  discussed  further  in  this  paper. 


Tribe  LABENINI 

Labenines  are  characterized  by  having  a  distinct  sclerotized  lobe  at  the  base  of  the  third  valvula. 
Most  species  also  have  fine  file-like  teeth  at  the  distal  apex  of  the  ovipositor.  The  tribe  contains 
four  genera:  Labena,  Certonotus,  Apechoneura  and  Xenothyris.  The  last  two  are  exclusively 
Neotropical,  and  Apechoneura  may  well  be  a  specialized  species-group  of  Certonotus.  Labena 
and  Certonotus  are  most  commonly  found  in  Australia  and  South  America,  and  there  is  evidence 
to  suggest  that  the  group  is  an  ancient  southern  one  that  originated  in  the  cool  temperate  forests 
of  Gondwanaland  (Gauld,  1983).  As  far  as  is  known,  the  native  hosts  of  the  group  are 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  111 

coleopterous  larvae  boring  in  wood  (Townes,  1969;  Gauld,  1984),  though  one  species  has 
adapted  to  parasitizing  introduced  siricids  (Hocking,  1967). 

Structurally,  labenines  are  amongst  the  most  primitive  ichneumonids.  Virtually  all  species 
have  clearly  developed  parapsidal  furrows,  a  primitive  hymenopterous  feature  found  widely 
amongst  Symphyta  and  some  lower  Aculeates.  An  apparently  unique  feature  of  the  group  is  the 
presence  of  a  distinct  sclerotized  lobe  at  the  base  of  the  third  valvula  (Fig.  4).  In  other 
ichneumonids  a  slightly  broadened,  rounded  area  is  present  in  a  homologous  position  and  this 
may  well  represent  a  precursor  of  the  valvular  lobe.  Quite  what  this  structure  is,  at  present  is 
unclear,  as  recent  morphological  treatises  make  no  mention  of  such  a  structure  (e.g.  Smith, 
1970;  Matsuda,  1976).  Short  (1978)  comments  on  the  primitive  structure  of  labenine  larvae. 

Key  to  genera  cf  Labenini  occurring  in  Australia 

1  Mesoscutum  punctate  or  puncto-striate;  occipital  carina  mediodorsally  complete  or  only 
narrowly  interrupted;  apex  of  fore  tibia  with  a  conspicuous  long  curved  spine  on  outer  distal 
margin;  female  with  third  fore  tarsal  segment  produced  apically  into  a  lobe  that  reaches 
beyond  apex  of  fourth  segment  (Fig.  7) LABENA  Cresson  (p.  Ill) 

-  Mesoscutum  with  transverse  rugae;  occipital  carina  dorsally  entirely  absent;  apex  of  fore  tibia 
with  a  short  tooth  on  outer  distal  margin;  female  with  fore  tarsus  simple 

CEtfrCWOritf  Kriechbaumer  (p.  117) 

LABENA  Cresson 

Labena  Cresson,  1864:  399.  Type-species:  Cryptus  grallator  Say,  by  subsequent  designation,  Viereck, 

1914:  80. 
Caryoecus  Walsh,  1866:  30.  Type-species:  Mesochorus  fuscipennis  Brulle  (=  Cryptus  grallator  Say),  by 

monotypy. 

Microtritus  Kriechbaumer,  1889:  307.  Type-species:  Microtritus  apicalis  Kriechbaumer,  by  monotypy. 
Dyseidopus  Kriechbaumer,  1890:  489.  Type-species:  Dyseidopus  sericeus  Kriechbaumer,  by  monotypy. 
Dysidopus  Schulz,  1906:  103.  [Unjustified  emendation.] 
Neonotus  Parrott,  1955:  230.  Type-species:  Neonotus  chadwickii  Parrott,  by  original  designation. 

Moderately  large  to  large  insects,  fore  wing  length  6-20  mm;  clypeus  small,  flat  or  concave,  very  thin  with 
margin  arcuate;  labrum  barely  projecting;  mandible  tapered,  twisted  30°  with  upper  tooth  slightly  the 
longer;  outer  mandibular  surface  with  a  median  longitudinal  groove  bearing  hairs;  malar  space  shorter 
than  basal  mandibular  width.  Occipital  carina  complete  or  narrowly  interrupted  centrally;  eye  margin 
slightly  indented  opposite  antennal  socket.  Antenna  almost  cylindrical,  that  of  female  with  a  small  flat 
sensillum  on  extreme  distal  apex.  Mesoscutum  punctate  or  striate  transversely,  with  notauli  vestigial; 
notaular  crests  well  developed.  Propodeum  rather  short,  convexly  rounded  with  spiracle  elliptical; 
propodeal  areae  usually  defined,  area  superomedia  usually  larger  than  area  petiolaris;  gaster  inserted  high 
on  propodeum,  above  level  of  hind  coxae.  Fore  tibia  of  female  inflated  (so  it  often  collapses  in  dried 
specimens)  with  a  large  curved  spine  on  outer  distal  margin;  fore  tarsus  with  segment  3  lobed,  the  lobe 
reaching  nearly  to  centre  of  segment  5,  segment  4  reduced;  hind  coxa  of  female  flattened  internally  with  a 
short  basal  groove;  tarsal  claws  large,  simple.  Fore  wing  with  cu-a  more  or  less  opposite  base  of  Rs&M; 
3r-m  present,  areolet  large,  rhombic  or  with  very  short  anterior  side;  2m-cu  sinuous  with  two  bullae.  Hind 
wing  with  distal  abscissa  of  Cu\  present,  sometimes  not  joining  to  first  abscissa  of  Cu\,  if  joined,  then  first 
abscissa  of  Cu\  is  shorter  than  cu-a;  basal  cell  slender;  Sc  with  one  or  two  hamuli.  Gaster  quite  long, 
cylindrical;  tergite  1  with  spiracles  a  little  before  centre;  sternite  1  usually  reaching  to  or  slightly  beyond 
level  of  spiracles;  laterotergites  2-4  membranous;  last  visible  tergite  with  a  small,  weakly  defined  dorsal 
plate  differentiated.  Ovipositor  projecting  beyond  apex  of  gaster  by  1-8-4-4  times  length  of  hind  tibia; 
upper  valve  with  apex  bearing  blunt  serrations,  lower  valve  apically  almost  enclosing  the  upper,  usually 
with  series  of  close  file-like  teeth  and  a  fine  coriaceous  patch  just  proximal  to  these  teeth,  rarely  with  teeth 
quite  strongly  developed. 

REMARKS.  Labena  is  a  large  genus  with  most  species  occurring  in  the  Neotropics.  Two  species  occur  in 
North  America,  and  in  this  work  seven  are  recognized  from  Australia. 

Labena  species  are  easily  separated  from  other  Labenini  by  the  specialized  fore  tarsus  and  the  possession 
of  a  sensillum  on  the  extreme  apex  of  the  antenna.  It  resembles  the  Neotropical  genus  Xenothyris  Townes 
in  having  a  relatively  smooth  mesoscutum,  though  in  the  latter  taxon  rugae  are  present  on  the  anterior  part 
of  the  median  lobe.  Unlike  Xenothyris,  Labena  species  have  simple  claws. 


112  I.  D.  GAULD  &  G.  A.  HOLLO  WAY 

The  relationships  of  the  Australian  species.  As  the  vast  majority  of  the  Neotropical  species  are 
undescribed  and  virtually  unstudied  it  is  impossible  as  yet  to  state  whether  the  Australian  Labena 
constitute  a  holophyletic  group,  so  no  formal  cladistic  analysis  is  presented  here.  However,  the  seven 
Australian  species  are  clearly  divisible  into  two  monophyletic  species-groups,  each  of  which  is  charac- 
terized by  several  apomorphic  features.  The  annulata-group  contains  four  taxa  (L.  annulata,  grandis, 
chadwickii  and  malecasta).  All  these  species  have  Cu\  in  the  hind  wing  incomplete  and  have  the  white 
flagellar  band  at  the  distal  apex  of  the  antenna.  The  keira-group  contains  three  taxa  (L.  keira,  jacunda  and 
pudenda).  These  species  have  virtually  a  smooth  mesopleuron,  and  the  male  genitalia  are  specialized  in 
that  the  apex  of  the  gonosquama  is  flattened  or  impressed  and  surrounded  by  a  fringe  of  very  long  hairs 
(Figs  9, 10). 

The  interrelationship  of  the  taxa  within  these  species-groups  is  less  clear.  Within  the  keira-group,  keira 
andpudenda  share  two  apparently  apomorphic  features,  a  specialized  mid  tibia  and  a  medially  interrupted 
yellow  band  on  the  central  female  gastral  segments.  The  unspecialized  mid  tibia  found  in  L.  jacunda  may 
represent  a  reversal  to  the  plesiomorphic  condition  unless  the  apomorphic  state  has  been  derived 
independently  in  both  the  keira  and  annulata  lineages  (and  probably  also  in  various  Neotropical  groups).  It 
is  therefore  arguable  that  the  possible  sister-relationship  between  keira  and  pudenda  is  only  supported  by 
the  colour  character.  Contradicting  this  relationship  are  two  features  shared  by  pudenda  and  jacunda  -  the 
coarse  ovipositor  teeth  (Fig.  21)  and  the  large  impressed  area  on  the  apex  of  the  male  gonosquama.  The 
latter  is  quite  clearly  an  apomorphic  condition  but  the  polarity  of  the  former  is  controversial  as  the 
possession  of  coarse  teeth  is  probably  plesiomorphic  for  ichneumonids.  Other  Labenini  all  have  fine, 
file-like  ovipositor  teeth  and  it  is  considered  that  this  condition  is  an  apomorphy  of  the  tribe  (Gauld,  1983) 
and  consequently  plesiomorphic  for  Labena  species.  Thus  the  possession  of  coarse  teeth  can  be  regarded  as 
a  further  apomorphic  development,  albeit  a  reversal,  supporting  the  group  pudenda  and  jacunda. 

Within  the  annulata-group  the  three  species  grandis,  chadwickii  and  malecasta  seem  to  form  a  subgroup 
characterized  by  possession  of  transverse  striations  on  the  mesoscutum.  This  subgroup  apparently  has  a 
sister-relationship  to  L.  annulata  (Fig.  65). 

Key  to  species  of  Labena  occurring  in  Australia 

1  Hind  wing  with  distal  abscissa  of  Cu\  present  in  membrane,  not  confluent  with  Cu\  and  cu-a  (Fig. 

11);  mesopleuron  quite  closely  punctate  or  punctocoriaceous,  often  submatt;  male  gonos- 
quama not  distally  strongly  flattened,  with  a  fringe  of  fine  hairs  (Fig.  8)  or  a  single  tuft  of  long 
hairs  ( a/inu/afa-group) 2 

-  Hind  wing  with  distal  abscissa  of  Cu^  complete  (Fig.  12);  mesopleuron  smooth  and  polished,  at 

most  with  sparse  punctures;  male  gonosquama  distally  flattened  or  impressed,  with  this  area 
fringed  with  long  hairs  (Figs  9, 10)  (Iteira-group) 5 

2  Tergites  2-4  of  gaster  coarsely  and  very  closely  punctate,  matt  (Fig.  13);  ovipositor  2-0  or  less 

times  as  long  as  hind  tibia annulata  (Brulle)  (p.  113) 

-  Tergites  2-4  of  gaster  smooth  and  polished ,  at  most  with  very  fine  punctures  (Fig .  14) ;  ovipositor 

at  least  2-5  times  as  long  as  hind  tibia 3 

3  Posterior  end  of  gaster  with  mediodorsal  margin  thickened  and  produced  into  a  rounded 

prominence  (Fig.  15);  pronotum  with  a  median  lateral  conical  tubercle;  metapleuron  strongly 
vermiculate grandis  sp.  n.  (p.  1 14) 

-  Posterior  end  of  gaster  with  mediodorsal  margin  thin,  slightly  concave,  not  produced  (Fig.  16); 

pronotum,  at  most,  with  a  median  lateral  convex  swelling,  usually  virtually  flat;  metapleuron 
punctate  or  puncto-striate 4 

4  Female  with  ovipositor  at  least  3-5  times  as  long  as  hind  tibia;  occipital  carina  mediodorsally 

complete;  male  with  a  tuft  of  long  hairs  on  distal  apex  of  gonosquama malecasta  sp.  n.  (p.  116) 

-  Female  with  ovipositor  at  most  3  •  0  times  as  long  as  hind  tibia ;  occipital  carina  (of  female  at  least) 

mediodorsally,  narrowly  interrupted;  male  with  gonosquama  distally  evenly  rounded  and 
uniformly  hirsute chadwickii  (Parrott)  (p.  1 13) 

5  Mid  tibia  slender,  subcylindrical  (Fig.  17);  tergites  4-5  of  gaster  with  broad  posterior  yellow 

marginal  band;  hind  coxa  of  female  with  ovipositor  guide  barely  differentiated 

jacunda  sp.  n.  (p.  115) 

-  Mid  tibia  distally  swollen,  proximally  narrowed  and  at  least  slightly  flattened  (Fig.  18);  tergites 

4-5  of  gaster  with  posterior  marginal  yellow  band  centrally  interrupted  or  yellow  band 
indistinct ;  hind  coxa  of  female  with  ovipositor  guide  well  developed 6 

6  Apex  of  fore  wing  infumate;  mid  tibia  with  a  longitudinal  row  of  3-6  spine-like  bristles  on  outer 

surface  (Fig.  18);  submetapleural  carina  evenly  and  weakly  broadened  anteriorly  (Fig.  19); 
ovipositor  about  3-0  times  as  long  as  hind  tibia keira  sp.  n.  (p.  115) 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  113 

-  Apex  of  fore  wing  very  uniformly  hyaline;  mid  tibia  without  obvious  spine-like  bristles  on  outer 
surface;  submetapleural  carina  anteriorly  abruptly  expanded  into  a  rounded  lobe  (Fig.  20); 
ovipositor  about  4-0  times  as  long  as  hind  tibia pudenda  sp.  n.(p.  117) 

Labena  annulata  (Brulle) 
(Figs  8, 11, 13) 

Ephialtes  annulatus  Brulle,  1846:  86.  Holotype  9,  TASMANIA  (MNHN)  [examined]. 
Labena  annulata  (Brulle)  Townes  etal. ,  1961:  112. 

Female.  Moderately  large  species,  fore  wing  length  7-13  mm.  Malar  space  0-4  times  as  long  as  basal 
mandibular  width.  Occipital  carina  mediodorsally  narrowly  interrupted,  ventrally  joining  hypostomal 
carina  and  continuing  to  mandibular  base  as  a  distinct  carina,  not  developed  into  a  flange.  Pronotum 
laterally  almost  flat;  mesothorax  laterally  and  ventrally  bearing  dense,  short,  white  pubescence;  mesoscu- 
tum  closely  and  coarsely  punctate;  horizontal  mesopleural  furrow  indistinct;  metapleuron  closely  and 
coarsely  punctate;  submetapleural  carina  anteriorly  expanded  into  a  rounded  lobe.  Propodeum  with 
anterior  transverse  carina  present  centrally  separating  area  superomedia  from  area  basalis.  Fore  wing  with 
3r-m  and  2r-m  converging,  widely  separated  at  junction  with  Rs.  Hind  wing  with  distal  abscissa  of  Cui 
present  in  membrane,  not  confluent  with  Cu\  and  cu-a.  Mid  tibia  with  a  longitudinal  row  of  spine-like 
bristles  on  outer  surface,  proximally  slender,  distally  swollen  with  a  trace  of  groove  in  inner  surface.  Gaster 
with  sternite  10-7  times  as  long  as  the  hind  coxa,  reaching  to  level  of  spiracle;  tergites  2-4  coarsely  and  very 
closely  punctate,  matt;  apex  of  terminal  plate  of  tergite  8  simple.  Ovipositor  projecting  beyond  apex  of 
gaster  by  1-8-2-0  times  length  of  hind  tibia,  the  apex  bearing  very  fine  teeth. 

Coloration.  As  for  L.  pudenda  although  some  specimens  with  fore  and  mid  legs  entirely  yellow. 

Male.  Similar  to  female  but  with  fore  wing  length  7-8  mm;  malar  space  0-4  times  basal  mandibular  width; 
gaster  with  sternite  1 0-7-0-8  times  as  long  as  hind  coxa.  Gonosquama  with  an  apical  tuft  of  fine,  long  hairs. 

REMARKS.  L.  annulata  is  one  of  the  most  distinctive  species  of  Australian  Labena  and  can  easily  be 
recognized  by  its  very  close,  coarse  punctured  microsculpture,  particularly  on  the  tergites  of  the  gaster. 
The  ovipositor  is  shorter  than  that  of  any  other  species.  This  species  is  widespread  in  Australia  though  it 
seems  to  be  most  common  in  the  south  and  west. 

HOST  RECORDS.  Cerambycidae:  Ur acanthus  strigosus  Pascoe  (NMV). 

MATERIAL  EXAMINED 

Holotype  $,  Tasmania:  no  further  data  (MNHN). 

Queensland:  1  $ ,  14  km  W.  by  N.  Hope  Vale  Mission  (15-16S  144-59E),  x.1980  (Cardale)  (ANIC).  New 
South  Wales:  1  $ ,  2  d",  Kinchega  NP,  ix.1980  (Rodd)  (AM).  Victoria:  1  $ ,  Baxter,  parasite  on  Uracanthus 
strigosus  (Dixon)  (NMV);  1  $,  Healesville,  xi.1983  (NMV).  Northern  Territory:  2  £,  Areyonga,  600  m, 
viii (TC).  Tasmania:  2  $,  Hobart, ix.1921  (Cole)  (NMV);  1  c?,  Ridgeway ix.1943  (Cole)  (NMV);  1  $,1867 
(Sichel)  (TC)  (compared  with  type).  Western  Australia:  1  $,  13  km  WNW.  Northampton  (28-18S 
114-31E),  x.1981  (Naumann  &  Cardale)  (ANIC);  1  $ ,  no  further  data  (BMNH). 

Labena  chadwickii  (Parrott) 
(Fig.  14) 

Neonotus  chadwickii  Parrott,  1955:  230.  Holotype  $,  NEW  SOUTH  WALES  (NSWDA)  [examined]. 
Labena  chadwickii  (Parrott)  Townes  &  Townes,  1960:  531. 

Female.  Moderate  to  large  species,  fore  wing  length  6-17  mm.  Malar  space  0-5  times  as  long  as  basal 
mandibular  width.  Occipital  carina  mediodorsally  narrowly  interrupted  at  least  in  female,  ventrally  joining 
hypostomal  carina  and  continuing  to  base  of  mandibular  base  as  distinct  carina,  not  developed  into  a 
flange.  Pronotum  laterally  with  a  median  swelling;  mesothorax  laterally  and  ventrally  bearing  sparse, 
short,  white  pubescence;  mesoscutum  punctate,  at  least  transversely  striate  in  centre,  in  larger  specimens 
uniformly  striate;  horizontal  mesopleural  furrow  distinct;  metapleuron  punctate;  metapleural  carina 
expanded  anteriorly  into  a  small  rounded  lobe.  Propodeum  with  anterior  transverse  carina  present 
centrally,  separating  area  superomedia  from  area  basalis.  Fore  wing  with  3r-m  and  2r-m  converging, 
almost  joining  at  junction  with  Rs.  Hind  wing  with  distal  abscissa  of  C«i  present  in  membrane,  not 
confluent  with  Cu\  and  cu-a.  Mid  tibia  with  a  longitudinal  row  of  spine-like  bristles  on  outer  surface, 
proximally  slender,  distally  swollen  and  with  indication  of  groove  on  inner  surface.  Gaster  with  sternite  1 
0-8  times  as  long  as  hind  coxa,  reaching  from  opposite  to  well  behind  level  of  spiracle;  tergites  2-4  smooth 


114  I.  D.  GAULD  &  G.  A.  HOLLOW  AY 

and  polished,  at  most  with  very  fine  punctures;  apex  of  terminal  plate  of  tergite  8  simple.  Ovipositor 
projecting  beyond  apex  of  gaster  by  2-2-3-0  times  length  of  hind  tibia,  the  apex  bearing  very  fine  teeth. 

Coloration.  Black.  Palps,  labrum,  clypeus,  face  (except  median  longitudinal  area),  genae,  orbits, 
anterolateral  margin  of  mesonotum,  tegula,  posterior  0-3  of  subalar  process,  posterior  0-5  of  scutellum, 
postscutellum,  posterior  0-3  of  propodeum,  posterior  margins  of  tergites  1-7,  distal  end  of  each  coxa,  fore 
and  mid  legs  with  trochanters,  distal  0-2  of  femur,  tibia  and  tarsus,  hind  legs  with  distal  0-1  of  femur  and 
tibial  spurs  yellow.  Median  area  of  frons,  mesonotum,  mesopleuron,  metapleuron,  upper  surface  of  hind 
coxa  partly  deep  red.  Antenna  with  disruptive  mark  subapical,  last  2-3  flagellar  segments  black. 

Male.  Similar  to  female  but  with  fore  wing  length  6-9  mm;  malar  space  0-6  times  basal  mandibular  width; 
gaster  with  sternite  1  0-8  times  as  long  as  hind  coxa.  Gonosquama  distally  evenly  rounded  and  uniformly 
hirsute.  Coloration  as  for  female  with  less  deep  red  areas. 

REMARKS.  L.  chadwickii  is  the  commonest  and  most  widespread  eastern  species  in  Australia.  Of  the  species 
with  an  apical  white  flagellar  band,  complete  yellow  posterior  margined  gastral  tergites  and  incomplete  Cu\ 
in  the  hind  wing  only  chadwickii  has  a  simple  terminal  plate,  smooth  gastral  tergites  and  incomplete 
occipital  carina.  The  females  are  rather  similar  to  L.  malecasta  but  differ  in  having  a  shorter  ovipositor  and 
the  genal  orbits  more  broadly  yellow  and  confluent  with  the  yellow  on  the  face.  L.  malecasta  has  only 
narrow  yellow  marks  on  the  genae  and  these  are  not  confluent  with  the  facial  yellow. 

HOST  RECORDS.  Buprestidae:  Ethon  affine  Laporte  &  Gory  (Parrott,  1955).  Several  individuals  have  been 
reared  in  Tasmania  from  logs  of  Pinus  radiata  infested  by  Sirex. 

MATERIAL  EXAMINED 

Holotype  $,  New  South  Wales:  Middlecove  Point,  Willoughby,  Sydney  (Chadwick)  (NSWDA). 

Queensland:  1  $,Eungella,xi(TC);  1  $,6mN.  of  Kuranda,  ll.i.1961  (McAlpine&Holloway)  (AM);  1 
$,  Mt  Cootha,  iv  (TC);  1  $,  Mt  Glorious,  vi.1977  (Miller)  (BMNH);  1  $,  Mt  Glorious,  xii.1976  (Boucek) 
(BMNH);  2  $ ,  Mt  Glorious,  i  (TC);  2  $ ,  Mt  Nebo,  ii  (TC);  1  $ ,  Mt  Tambourine,  iii.1950  (Burns)  (NMV); 
4  $,  Mt  Tambourine,  xii  (TC);  1  $,  O'Reilly's  Guest  House,  iii.1980  (BMNH);  1  cf,  Palm  Is.,  near 
Townsville,  x  (TC);  1  cf,  Stanthorpe,  1982  (Sedlacek)  (TC).  New  South  Wales:  1  $,  Eucambene  Dam, 
i.1961  (Leipa)  (ANIC);  3  cf  (paratypes),  same  data  as  holotype  (AM,  ANIC,  NSWDA);  1  $,  Monga, 
ix.1957  (ANIC);  6  cf,  Mt  Toman,  ix.1982  (Rodd)  (AM);  1  $,  Mt  Tomah,  xi.1978  (Rodd)  (AM);  1  $, 
Wentworth  Falls,  Blue  Mtns,  xi.1982  (McAlpine)  (AM).  Australian  Capital  Territory:  1  <j>,  Lees  Ck, 
xi.1977  (Daniels)  (AM).  Victoria:  1  $ ,  6  km  S.  Aberfeldy,  xi.1976  (Colder)  (NMV);  1  $ ,  Blackwood  Ra., 
x.1953  (Neboiss)  (NMV);  2  $,  Ferntree  Gully,  x.1951  (Oke)  (NMV);  1  $>,  Gippsland  (AM);  1  cf, 
Kinglake  W.,  x.1954  (Neboiss)  (NMV);  1  $,  Lai  Lai,  i.1954  (Neboiss)  (NMV);  1  $,  1  cf,  12  km  SE. 
Merrijig,  Howqua  R.,  xi.1971  (Neboiss)  (NMV);  1  $,  Mitta  Mitta  R.,  x.1973  (NMV);  1  $,  1  cf,  Mt 
Dandenong,  200  m,  ii  (TC);  1  cf,  Mt  Drummer,  xii. 1956  (Riek)  (ANIC);  1  $,  Warburton,  xii. 1972 
(Sedlacek)  (TC);  1  $,  Woori  Yallock,  x.1932  (Burns)  (NMV).  Tasmania:  1  cf ,  Bronte  Park,  i  (TC);  1  $, 
Cambridge,  i.1965,  ex  Pinus  radiata  log  (Taylor)  (ANIC);  1  $,  Cambridge,  xii. 1964,  exPinus  radiata  logs 
(Taylor)  (BMNH);  1  $,  2  cf ,  Hobart,  xi.1922  (Cole)  (NMV);  1  $,  Hobart,  xii.1915  (Cole)  (NMV);  2  cf , 
Launceston,  x.1927  (Cole)  (NMV);  1  $,  Leven  R.,  xi.1975  (Neboiss);  1  $,  Ridgeway,  ix.1942  (Cole) 
(NMV);  2  $,  Roseberry,  i  (TC);  1  $,  Strahan,  iii  (TC). 

Labenagrandissp.  n. 

(Fig.  15) 

Female.  Large  species,  fore  wing  length  17  mm.  Malar  space  0-6  times  as  long  as  basal  mandibular  width. 
Occipital  carina  mediodorsally  complete,  ventrally  joining  hypostomal  carina  and  continuing  to  mandibu- 
lar base  as  distinct  carina,  not  forming  a  flange.  Pronotum  laterally  with  a  median  conical  tubercle; 
mesothorax  laterally  and  ventrally  bearing  sparse,  short,  white  pubescence;  mesoscutum  strongly, 
transversely  striate;  horizontal  mesopleural  furrow  distinct,  at  least  posteriorly;  metapleuron  vermiculate; 
submetapleural  carina  strongly  broadened  into  a  triangular  striate  lobe.  Propodeum  with  anterior 
transverse  carina  present  centrally,  separating  area  superomedia  and  area  basalis.  Fore  wing  with  2r-m 
converging  towards  3r-m,  almost  joining  it  at  junction  with  Rs.  Hind  wing  with  distal  abscissa  of  Cu\ 
present  on  membrane,  not  confluent  with  Cu\  and  cu-a.  Mid  tibia  with  longitudinal  row  of  spine-like 
bristles  on  outer  surface,  the  distal  end  swollen  somewhat,  proximally  slender  and  with  a  diagonal  internal 
furrow  present.  Gaster  with  sternite  10-5  times  as  long  as  hind  coxa,  reaching  to  level  of  spiracle;  tergites 
2-4  smooth  and  polished ;  apex  of  terminal  plate  of  tergite  8  specialized.  Ovipositor  projecting  beyond  apex 
of  gaster  by  2-8  times  length  of  hind  tibia,  the  apex  bearing  very  fine  teeth. 

Coloration.  Black.  Palps,  labrum,  clypeus,  inner  orbits,  outer  orbits,  antero-lateral  margin  of  mesoscu- 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  115 

turn ,  tegula ,  posterior  0  •  5  of  subalar  process ,  posterior  0  •  4  of  propodeum ,  posterior  margin  of  tergites  1-7 , 
fore  and  mid  legs  (except  inner  surface  of  femur  and  tibia),  hind  leg  with  distal  tip  of  coxa,  trochanter, 
femur,  proximal  end  of  tibia  and  tarsus  yellow. 
Male.  Unknown. 

REMARKS.  Only  a  single  specimen  of  this  species  is  known.  It  is  immediately  distinguished  by  possession  of  a 
uniquely  specialized  tergite  8,  having  the  metapleuron  strongly  vermiculate  and  having  a  swollen 
pronotum.  The  hind  coxae  are  shorter  and  stouter  than  those  of  other  species  and  the  petiole  is 
considerably  more  robust. 

MATERIAL  EXAMINED 
Holotype  $,  Western  Australia:  Yallingup,  Cape  Naturaliste,  ix-x.1913  (Turner)  (BMNH). 

Labenajacunda  sp.  n. 

(Fig.  17) 

Female.  Large  species,  fore  wing  length  13-14  mm.  Malar  space  0-6  times  as  long  as  basal  mandibular 
width.  Occipital  carina  mediodorsally  complete,  ventrally  joining  hypostomal  carina  far  from  base  of 
mandible.  Pronotum  laterally  weakly  convex;  mesothorax  laterally  and  ventrally  bearing  sparse  pubes- 
cence; mesoscutum  with  superficial  punctures;  horizontal  mesopleural  furrow  indistinct;  metapleuron 
finely  sparsely  punctate;  submetapleural  carina  broadened  anteriorly  into  ribbed  lobe.  Propodeum  with 
anterior  transverse  carina  complete  so  area  superomedia  is  distinctly  delineated.  Fore  wing  with  3r-m 
slightly  inclined,  2r-m  strongly  convergent  anteriorly.  Hind  wing  with  distal  abscissa  of  Cui  complete.  Mid 
tibia  slender,  cylindrical,  with  scattered  spines  on  outer  surface.  Gaster  with  sternite  1 0-6-0-7  times  as  long 
as  hind  coxa,  reaching  to  level  of  spiracle,  the  membranous  portion  unusual  in  being  a  pair  of  well- 
developed  crests;  tergites  2-4  smooth  and  polished;  apex  of  terminal  plate  of  tergite  8  simple.  Ovipositor 
projecting  beyond  apex  of  gaster  by  about  3-5  times  length  of  hind  tibia,  apex  bearing  8  distinct  strong 
teeth. 

Coloration.  Black.  Face,  frontal  and  genal  orbits  narrowly,  tegula,  small  mark  on  subalar  prominence, 
scutellum,  posterior  part  of  propodeum  and  posterior  margins  of  all  tergites  yellow;  mesopleuron  and  base 
of  petiole  reddish;  anterior  two  pairs  of  legs  predominantly  yellow;  hind  legs  black,  distal  apex  of  coxa  and 
femur  and  proximal  end  of  tibia  and  basitarsus  yellow. 

Male.  Similar  to  female  but  with  fore  wing  length  13-14  mm;  malar  space  0-3  times  basal  mandibular 
width;  gaster  with  sternite  11-0  times  as  long  as  hind  coxa.  Gonosquama  distally  flattened,  impressed,  the 
impressed  area  surrounded  by  a  fringe  of  long  hairs.  Black  with  face,  frontal  and  genal  orbit,  tegula, 
subalar  prominence,  scutellum,  postscutellum,  hind  margin  of  propodeum,  broad  posterior  bands  on 
gastral  tergites,  anterior  2  pairs  of  legs,  hind  trochanter  and  trochantellus,  proximal  0-5  of  tibia,  entire 
basitarsus  and  extreme  proximal  apex  of  second  tarsal  segment  yellow.  Flagellum  black,  with  a  subapical 
whitish  mark  that  is  not  a  complete  band. 

REMARKS.  The  female  of  this  species  is  immediately  recognizable  by  the  slender  mid  tibia  and  ventral 
petiolar  prominences  which  are  unique  features  of  this  species  amongst  Australian  Labena.  The  broad 
yellow  bands  on  the  gaster  distinguish  the  male  ofjacunda  from  pudenda  and  keira,  the  only  other  species 
with  ornamented  male  gonosquamae. 

MATERIAL  EXAMINED 
Holotype  $,  Victoria:  Chiltern  (NMV). 
Paratypes.  Victoria:  2  cT,  Mt  Buffalo,  4,500',  13.U955  (Neboiss)  (NMV). 

Labena  keira  sp.  n. 

(Figs  9, 12, 18, 19) 

Female.  Large  species,  fore  wing  length  11-16  mm.  Malar  space  0-5  times  as  long  as  basal  mandibular 
width.  Occipital  carina  mediodorsally  complete,  ventrally  joining  hypostomal  carina  and  forming  a  flange. 
Pronotum  laterally  with  a  slight  median  swelling;  mesothorax  laterally  and  ventrally  bearing  no  white 
pubescence;  mesoscutum  smooth,  with  sparse  fine  punctures;  horizontal  mesopleural  furrow  distinct  and 
widening  posteriorly;  metapleuron  smooth  and  polished;  submetapleural  carina  slightly  broadened 
anteriorly,  not  produced  into  a  lobe.  Propodeum  with  anterior  transverse  carina  present  centrally, 
separating  area  superomedia  from  area  basalis.  Fore  wing  with  3r-m  and  2r-m  convergent,  sometimes 
almost  joining  at  junction  with  Rs.  Hind  wing  with  distal  abscissa  of  C«i  complete.  Mid  tibia  with  a 
longitudinal  row  of  spine-like  bristles  on  outer  surface,  proximally  slender  and  slightly  flattened,  distally 


116  I.  D.  GAULD  &  G.  A.  HOLLOWAY 

swollen.  Gaster  with  sternite  1  0-7  times  as  long  as  hind  coxa,  just  reaching  to  slightly  behind  level  of 
spiracle ;  tergites  2-4  smooth  and  polished ;  apex  of  terminal  plate  of  tergite  8  simple .  Ovipositor  projecting 
beyond  apex  of  gaster  by  3  times  length  of  hind  tibia,  the  apex  bearing  very  fine  teeth. 

Coloration.  Black  to  deep  red.  Palps,  labrum,  clypeus,  face,  outer  orbits,  frons  (except  ocellar  triangle 
and  vertex),  scutellum,  scutellar  ridges,  postscutellum,  postscutellar  ridges,  tegulae,  subalar  process,  small 
mesopleural  macula,  posterior  0-5  of  propodeum,  posterior  margin  of  tergites  1-8  (often  divided 
medially),  fore  and  mid  legs  except  sometimes  distal  0-5  of  mid  femur,  hind  trochanters,  basal  0-5  of  tibia 
and  basal  tarsal  segment  yellow.  Disruptive  mark  of  antenna  medially  placed,  apex  of  fore  wing  infumate. 

Male.  Similar  to  female  but  with  fore  wing  length  8-13  mm;  malar  space  0-4  times  basal  mandibular 
width;  gaster  with  sternite  1  0-9  times  as  long  as  hind  coxa.  Gonosquama  distally  flattened  and  fringed 
partially  by  very  fine  hairs.  Coloration.  As  for  female,  sometimes  with  less  yellow  on  gaster  and  no  white 
disruptive  marks  on  antenna. 

REMARKS.  L.  keira  is  immediately  recognizable  by  the  infumate  mark  on  the  distal  apex  of  the  fore  wing. 
Structurally  it  is  rather  similar  to  L.  pudenda  from  which  it  may  be  separated  by  the  submetapleural  carina 
being  barely  expanded,  spinose  mid  tibia  and  centrally  complete  anterior  transverse  carina  of  the 
propodeum.  This  is  one  of  the  most  widely  distributed  species  of  Labena  and  is  recorded  from  Victoria 
north  to  central  Queensland.  It  is  also  known  to  occur  on  Lord  Howe  Island. 

MATERIAL  EXAMINED 

Holotype  $,  New  South  Wales:  Mt  Tomah,  28.iii.1980  (Rodd)  (AM). 

Paratypes.  Queensland:  1  $,  Bunya  Mtns,  i.1938  (Geary}  (AM);  3  cf ,  Eungella,  ix.1923  (NMV);  2  $, 
Eungella,  xi  (TC);  1  $ ,  9  cf ,  Montville,  ix.1955  (Bums)  (NMV);  1  $ ,  9  cf ,  Mt  Glorious,  xi  &  i  (TC);  4  $ ,  1 
Cf ,  Mt  Tambourine,  x.  1977  (Galloway)  (BMNH);  7  $ ,  8  cf,  Mt  Tambourine,  xi-xii  (TC);  1  $ ,  Mt  Tip  Tree 
(17-02S  145-37E),  x.1980  (Cardale)  (ANIC);  1  cf,  Westwood,  xi.1924  (Burns)  (NMV);  1  cf ,  Wilson's 
Peak,  Killarney,  9.U977  (Boucek)  (BMNH).  New  South  Wales:  1  cf ,  Lord  Howe  Is.,  (AM);  1  cf ,  Lord 
Howe  Is.,  30.xi.1955  (Paramonov  &  Leipa)  (ANIC);  1  cf ,  Lord  Howe  Is.  xii.1977  (Liepa)  (ANIC);  1  cf , 
Mooney  Mooney  Creek,  near  Gosford,  xi.1975  (McAlpine  &  Schneider)  (AM).  Australian  Capital 
Territory:  1  $,  Mt  Gingera,  i.1957  (Riek)  (ANIC). 

Labena  malecasta  sp.  n. 

(Fig.  16) 

Female.  Large  species,  fore  wing  length  13-15  mm.  Malar  space  0-8  times  as  long  as  basal  mandibular 
width.  Occipital  carina  mediodorsally  complete,  ventrally  joining  hypostomal  carina  and  continuing  to 
mandibular  base  as  a  distinct  carina,  not  developed  into  a  flange.  Pronotum  laterally  with  a  median 
swelling;  mesothorax  laterally  and  ventrally  bearing  sparse,  long,  white  pubescence;  mesoscutum  finely 
punctate  with  indications  of  weak  transverse  striae;  horizontal  mesopleural  furrow  indistinct;  metapleuron 
indistinctly  puncto-striate;  submetapleural  carina  anteriorly  abruptly  expanded  into  an  almost  quadrate 
lobe.  Propodeum  with  anterior  transverse  carina  present  centrally,  separating  area  superomedia  from  area 
basalis,  latter  with  transverse  striae.  Fore  wing  with  3r-m  converging  towards  2r-m,  almost  joining  at 
junction  with  Rs.  Hind  wing  with  distal  abscissa  of  C«i  present  in  membrane,  not  confluent  with  Cu\  and 
cu-a.  Mid  tibia  with  a  longitudinal  row  of  spine-like  bristles  on  outer  surface,  proximally  slender,  distally 
swollen,  with  an  oblique  furrow  on  inner  surface.  Gaster  with  sternite  1  0-7  times  as  long  as  hind  coxa, 
reaching  to  well  behind  level  of  spiracle;  tergites  2-4  smooth  and  polished  with  sparse,  fine  punctures;  apex 
of  terminal  plate  of  tergite  8  simple .  Ovipositor  pro j  ecting  beyond  apex  of  gaster  by  at  least  3  •  5  times  length 
of  hind  tibia,  the  apex  bearing  distinct  teeth. 

Coloration.  Black  to  red.  Palps,  clypeus,  inner  orbit  of  face,  emargination  of  orbit  opposite  antennal 
socket,  thin  line  on  outer  orbit,  antero-lateral  corner  of  mesoscutum,  tegula,  subalar  process,  posterior  0-3 
of  scutellum,  postscutellum,  posterior  0-3  of  propodeum,  posterior  margin  of  tergites  1-7,  fore  leg  (except 
coxa  and  distal  part  of  femur),  mid  leg  with  distal  end  of  femur,  tibia  and  tarsal  segments  1-4,  hind  leg  with 
distal  end  of  tibia  and  basal  tarsal  segment  yellow.  Antennal  disruptive  mark  at  distal  end  except  for  last 
1-2  segments  which  are  blackish. 

Male.  Similar  to  female  but  with  fore  wing  length  12  mm;  malar  space  0-5  times  basal  mandibular  width; 
gaster  with  sternite  10-9  times  as  long  as  hind  coxa.  Gonosquama  with  an  apical  tuft  of  long  hairs. 
Coloration  similar  to  female. 

REMARKS.  This  large  species  is  structurally  rather  similar  to  L.  chadwickii.  The  female  has  a  subtly  different 
colour  pattern  and  a  longer  ovipositor  whilst  the  male  is  distinct  in  bearing  a  tuft  of  long  hair  on  the 
gonosquama.  L.  malecasta  is  only  known  from  Tasmania,  Victoria  and  the  southern  alps  of  New  South 
Wales. 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  117 

MATERIAL  EXAMINED 

Holotype  $,  Tasmania  (BMNH). 

Paratypes.  New  South  Wales:  1  cf,  Mt  Kosciusko,  Dainer's  Gap,  x.1929  (Musgrave)  (AM).  Victoria: 
1  9,  Warburton,  ii-iii  (TC).  Tasmania:  1  $  (BMNH). 

Labena  pudenda  sp.  n. 

(Figs  10,  20,  21) 

Female.  Large  to  very  large  species,  fore  wing  length  13-20  mm.  Malar  space  0-7  times  as  long  as  basal 
mandibular  width.  Occipital  carina  mediodorsally  complete,  ventrally  joining  hypostomal  carina  and 
forming  flange.  Pronotum  laterally  with  a  median  convex  swelling;  mesothorax  laterally  and  ventrally 
bearing  sparse,  short  white  pubescence;  mesoscutum  smooth  with  sparse  fine  punctures;  horizontal 
mesopleural  furrow  distinct;  mesopleuron  smooth  and  polished;  metapleuron  smooth  and  polished; 
submetapleural  carina  anteriorly  expanded  with  a  large  rounded  striate  lobe.  Propodeum  with  anterior 
transverse  carina  absent  centrally  so  areae  superomedia  and  basalis  are  confluent.  Fore  wing  with  3r-m 
slightly  inclined,  2r-m  strongly  converging  towards  3r-m,  both  widely  separated  at  junction  with  Rs.  Hind 
wing  with  distal  abscissa  of  C«i  complete.  Mid  tibia  without  obvious  spine-like  bristles  on  outer  surface, 
proximally  slender  and  slightly  flattened,  distally  swollen.  Gaster  with  sternite  1  0-7-1-0  times  as  long  as 
hind  coxa,  reaching  behind  level  of  spiracle;  tergites  2-4  smooth  and  polished;  apex  of  terminal  plate  of 
tergite  8  simple.  Ovipositor  projecting  beyond  apex  of  gaster  by  4-0-4-4  times  length  of  hind  tibia,  the  apex 
bearing  strong  teeth. 

Coloration.  Black  to  deep  red.  Palps,  labrum,  clypeus,  inner  and  outer  orbits,  ventral  corner  of 
propleuron,  tegula,  subalar  process,  scutellum,  postscutellum,  posterolateral  corner  of  propodeum, 
posterior  margin  of  tergites  1-7  (may  be  divided  medially),  fore  and  mid  leg  with  distal  part  of  coxa, 
trochanter,  femur,  distal  and  proximal  ends  of  tibia  and  tarsal  segments  1-4,  hind  leg  with  distal  end  of  each 
segment  including  tarsal  segment  1  and  entire  tarsal  segments  2-4  yellow. 

Male.  Similar  to  female  but  with  fore  wing  length  10-11  mm;  malar  space  0-9  times  basal  mandibular 
width;  gaster  with  sternite  1  0-9  times  as  long  as  hind  coxa.  Gonosquama  distally  flattened  and  fringed 
almost  entirely  by  very  fine  hairs.  Coloration  similar  to  female. 

REMARKS.  This  large,  rather  slender  species  is  easily  recognized  by  the  very  long  ovipositor  of  the  female. 
The  tip  of  this  organ  is  distinctive  in  having  much  coarser  teeth  than  are  usually  found  in  species  of  Labena. 
Males  have  the  most  specialized  gonosquamae  of  any  Australian  Labena  species.  L.  pudenda  seems  to  be  a 
southern  temperate  forest  species  and  has  been  recorded  from  New  South  Wales,  Victoria  and  the 
Australian  Capital  Territory. 

MATERIAL  EXAMINED 

Holotype  $,  Victoria:  BogongHigh  Plains,  xii.1931  (Kubala)  (NMV). 

Paratypes.  New  South  Wales:  1  cf,  Dainer's  Gap  (36-12S  148-43E),  xi.1973  (Morrow)  (ANIC);  1  $, 
Kosciusko,  xii.1922  (Goldfinch)  (AM);  1  $,  Mt  York,  x.1930  (NMV).  Australian  Capital  Territory:  1  $, 
Lees  Springs,  xi.1953  (Riek)  (ANIC).  Victoria:  1  ? ,  1  cf ,  'Alps',  xii.  1910  (NMV);  1  $ ,  Mt  Buffalo,  1600  m, 
ii  (TC);  1  ? ,  2  cf ,  'Victoria',  ii.1901  (French)  (BMNH);  1  $ ,  Yarra  Falls,  S.  Warburton,  i.1907  (Barnard) 
(NMV). 

CERTONOTUS  Kriechbaumer 

Certonotus  Kriechbaumer,  1889:  308.  Type-species:  Certonotus  varius  Kriechbaumer,  by  monotypy. 
Asperellus  Townes  in  Townes  et  al.,  1961:  471.  Type-species:  Certonotus  hinnuleus  Krieger,  by  original 
designation.  Syn.  n. 

Small  to  very  large  insects,  fore  wing  length  4-17  mm;  clypeus  flat,  transverse,  margin  thin,  evenly  arcuate; 
labrum  barely  projecting;  mandible  short,  stout  but  tapered,  twisted  25-30°,  with  upper  tooth  the  longer; 
outer  mandibular  surface  with  a  groove  bearing  hairs;  malar  space  trans-striate,  usually  a  little  longer  than 
basal  mandibular  width.  Occipital  carina  dorsally  absent;  eye  not  indented  next  to  antennal  socket. 
Antenna  slightly  clavate,  apically  pointed,  without  a  flat  sensillum.  Mesoscutum  with  transverse  rugae, 
notauli  weak,  notaular  crests  very  weak.  Propodeum  usually  quite  short,  convexly  rounded  with  spiracle 
elliptical;  anterior  transverse  carina  usually  complete  except  centrally,  other  carinae  often  reduced,  area 
superomedia  usually  not  delineated;  gaster  inserted  high  up  on  propodeum,  above  level  of  hind  coxae. 
Fore  tibia  with  a  short  tooth  on  outer  distal  margin;  fore  tarsus  unspecialized;  hind  coxa  of  female  with  an 
anterior  carina  continued  ventrally  as  a  process,  the  area  behind  this  carina  concave  and  closely  punctate; 


118  I.  D.  GAULD  &  G.  A.  HOLLOW  AY 

tarsal  claws  simple.  Fore  wing  with  cu-a  opposite  or  proximal  to  base  of  Rs&M;  3r-m  usually  present, 
areolet  almost  triangular,  often  petiolate  above;  2m-cu  sinuous,  with  two  close  bullae.  Hind  wing  with 
distal  abscissa  of  Cu\  present  or  incomplete  or  absent;  first  abscissa  of  Cu\  shorter  than  cu-a;  basal  cell 
slender;  Sc  with  one  or  two  hamuli.  Gaster  quite  long,  tergite  1  from  stout  to  quite  slender,  with  spiracles 
before  centre;  sternite  1  usually  reaching  nearly  to  level  of  spiracles;  laterotergites  2-4  membranous, 
folded  under;  tergite  8  highly  modified,  projecting  laterally  as  a  pair  of  prominences  at  either  side  of 
ovipositor  base,  dorsally  with  a  detached  plate  projecting  through  concave  orifice  in  hind  margin,  tergite  7 
often  mediodorsally  incised.  Ovipositor  projecting  beyond  apex  of  gaster  by  3-0-8-0  times  length  of  hind 
tibia,  its  apex  compressed,  the  upper  valve  with  weak  blunt  serrations,  the  lower  valve  enclosing  the  upper 
and  bearing  fine  file-like  teeth. 

REMARKS.  Certonotus  is  a  large  genus  centred  in  Australia  with  a  few  species  present  in  New  Guinea,  New 
Zealand  and  South  America.  Previously,  the  species  now  included  in  this  taxon  were  divided  between 
Certonotus  and  Asperellus  (Townes,  1969;  Gauld,  1984)  but  more  detailed  study  has  revealed  that  the 
latter  genus  is  almost  certainly  polyphyletic.  In  the  present  work  23  species  are  recognized  as  occurring  in 
Australia. 

Certotonus  species  are  easily  recognized  by  the  possession  of  transverse  rugae  on  the  mesoscutum,  a 
feature  immediately  distinguishing  the  genus  from  all  other  Australian  labenines.  Certonotus  species  may 
inadvertently  be  confused  with  Rhyssini,  especially  Epirhyssa  species.  Unlike  rhyssines,  which  have  an 
undeveloped  submetapleural  carina,  Certonotus  has  a  broad  expanded  lobe  present  anteriorly.  Further- 
more, Certonotus  has  fine  file-like  teeth  on  the  ovipositor  apex,  not  the  coarse  teeth  found  in  Epirhyssa 
(Gauld,  1984). 

The  relationships  of  the  Australian  species.  Gauld  (1984)  suggested  that  Asperellus,  as  defined  by  Townes 
(1969),  was  merely  a  specialized  species-group  of  Certonotus.  Further  study  suggests  that  Asperellus  is  not 
even  a  monophyletic  group,  but  rather  an  assemblage  of  species  of  Certonotus  that  lack  the  distal  abscissa 
of  Cu\.  The  majority  of  species  (farrugiai,  pineus,  mogimbensis,  zebrus,  toolangi,  hinnuleus  and 
leeuwinensis)  do  comprise  a  natural  group,  the  leeuwinensis-group.  All  have  rather  similar  propodeal 
carination,  in  that  the  posterior  transverse  carina  is  absent  but  a  large  smooth  area  is  enclosed  by  the 
anterior  carina  and  the  lateral  carinae;  they  possess  a  very  small  areolet,  have  one  or  more  spine-like 
bristles  present  on  the  hind  tibia  and  have  a  slightly  convex  face.  Most  have  a  rather  short  first  sternite,  a 
quite  deeply  divided  tergite  7  and  a  pronounced  lobe  on  tergite  8.  Only  in  farrugiai  is  tergite  7  barely 
indented  posteriorly.  The  species  ixion,  paluma,  celeus  and  Certonotus  species  A  would  all  run  to 
Asperellus  in  Townes'  (1969)  key  but  these  do  not  appear  to  be  closely  related  to  the  others  (i.e.  the 
leeuwinensis-group).  The  first  three  are  closely  related  to  Certonotus  talus  and  belong  to  the  humeralifer- 
group,  which  is  defined  by  having  a  very  deeply  divided  tergite  7  and  possessing  rather  long  narrow 
processes  on  tergite  8.  The  group  includes  seven  Australian  species  -  humeralifer,  apicalis,  talus,  cestus, 
ixion,  paluma  and  Certonotus  species  A.  The  New  Zealand  species  C.  fractinervis  apparently  also  belongs 
to  this  group. 

The  majority  of  other  Australian  species  (annulatus,  nitidulus,  geniculatus,  rufescens,  andrewi,  avitus, 
sisyphus  and  celeus)  constitute  a  third  group,  having  a  weakly  to  moderately  deeply  divided  tergite  7, 
rounded  lobes  on  tergite  8  and  a  relatively  long  first  sternite.  Within  the  nitidulus-group  one  rather 
distinctive  lineage  can  be  recognized  comprising  rufescens,  geniculatus,  sisyphus  and  celeus.  These  species 
all  have  flat  lower  faces,  and  most  have  the  posterior  end  of  the  lateral  propodeal  carina  broadened  to  form 
a  raised  keel. 

The  remaining  Australian  species,  C.  monticola,  belongs  to  the/Zav/ce/w-group  which  is  characterized  by 
possession  of  elongate  glossae  and  a  very  short  occipital  carinal  stub.  Males  of  species  in  this  group  are 
specialized  in  having  a  very  elongate  gaster  (with  tergite  7  about  1-5  times  as  long  as  broad  or  longer). 
Frequently  the  hind  margins  of  the  tergites  are  concave.  This  elongate  form  of  the  gaster  is  characteristic  of 
wood-boring  species  in  which  the  male  copulates  with  the  female  prior  to  her  emergence  from  the  burrow 
(see  Nuttall,  1973) .  The  male  of  C.  fractinervis  resembles  males  of  theflaviceps-group  though  the  structure 
of  the  female  suggests  the  species  is  best  placed  in  the  humeralifer-group. 

The  relationships  of  the  species-groups  are  difficult  to  determine  as  there  is  considerable  conflict  in  the 
characters.  The  leeuwinensis-group,  a  holophyletic  clade,  may  well  be  the  sister-lineage  of  the  humeralifer- 
group.  Both  have  a  similarly  deeply  divided  tergite  7  and  many  often  possess  spine-like  bristles  on  the  hind 
tibia  and  show  reduction  in  the  distal  abcissa  of  Cu\.  This  arrangement  leads  to  difficulty  in  placing  C. 
farrugiai  which  has  tergite  7  barely  divided.  The  humeralifer-group  could  be  paraphyletic  with  respect  to 
the  leeuwinensis-group.  The  nitidulus-group  could  well  be  a  paraphyletic  assemblage,  the  stem  group  from 
which  all  others  have  arisen,  although  the  rufescens-subgroup  is  clearly  a  holophyletic  clade.  The 
flaviceps-group  is  also  undoubtedly  a  holophyletic  clade. 

The  geographical  distribution  of  the  species- groups  is  as  follows: 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  119 

nitidulus-group:  New  Guinea,  Australia; 
rufescens-subgroup:  Australia; 
humeralifer-group:  Australia,  New  Zealand; 
flaviceps-group:  Moluccas,  New  Guinea,  tropical  Australia; 
leeuwinensis-group:  New  Guinea,  Australia,  New  Hebrides,  New  Caldeonia. 

Key  to  species  of  Certonotus  occurring  in  Australia 

1  Hind  wing  with  distal  abscissa  of  Cui  complete 2 

-  Hind  wing  with  distal  abscissa  of  Cui  either  incomplete,  that  is  present  in  membrane  but  not 

joining  Ci/i  &  cu-a,  or  entirely  absent 13 

2  Gaster  more  or  less  entirely  yellowish  or  orange-brown ,  the  tergites  in  dorsal  view ,  unicolorous 

or  slightly  infurcate  along  posterior  margin 3 

-  Gaster  dark  reddish  brown  or  black,  usually  with  conspicuous  yellow  spots  or  bands  along 

margin  of  tergites ,  some  marks  with  gaster  unicolorous  dark  brown 7 

3  Subalar  prominence,  in  dorsal  view,  strongly  raised,  with  a  blunt,  back-curved,  thorn-like 

protuberance  which  is  more  sharply  pointed  in  the  male  than  the  female  (Fig.  22);  malar 
space  long  (Fig.  25),  in  female  0-9-1-0  times  as  long  as  basal  mandibular  width,  in  male 
0-8-0-9  times  as  long;  submetapleural  carinal  flange  with  a  ridge  delimiting  an  anterolateral 
triangular  area rufescens  Morley  (p.  133) 

-  Subalar  prominence  in  dorsal  view  at  most  convex,  not  produced  into  a  spine-like  protuberance 

(Figs  23,  24);  malar  space  fairly  short  (Fig.  26),  in  female  0-7-0-8  or  male  0-4-0-6  times  as 
long  as  basal  mandibular  width;  submetapleural  carinal  flange  without  a  delimited  triangular 
area  anterolaterally 4 

4  Pronotum  in  dorsal  view  with  part  before  upper  corner  strongly  convex  or  pyramidal  (Figs  23, 

24);  anterior  transverse  carina  of  propodeum  present  centrally  (Fig.  27);  2r-m  and  3r-m  fused 
anteriorly  so  areolet  is  petiolate  (Fig.  30) 5 

-  Pronotum  in  dorsal  view  with  part  before  upper  corner  flat  to  weakly  convex;  anterior 

transverse  carina  of  propodeum  absent  centrally  (Figs  28,  29);  2r-m  and  3r-m  joining  Rs 
separately  so  areolet  not  petiolate  (Fig.  31) 6 

5  Upper  part  of  pronotum  pyramidal  in  dorsal  view  (Fig.  24);  female  with  ovipositor  about  5 

times  as  long  as  hind  tibia;  hind  tibia  with  one  spine-like  bristle  on  posterior  margin; 
flagellum  entirely  black humeralifer  Krieger  (p.  127) 

-  Upper  part  of  pronotum  simply  strongly  convex  in  dorsal  view  (Fig.  23) ;  female  with  ovipositor 

7-5  or  more  times  as  long  as  hind  tibia;  hind  tibia  without  a  spine-like  bristle  or  posterior 
margin;  flagellum  with  distal  end  white apicalis  Morley  (p.  122) 

6  Posterior  transverse  carina  of  propodeum  centrally  strongly  raised  (Fig.  28);  antenna  with 

apical  10  or  so  segments  whitish;  spiracular  area  bounded  posteriorly  by  carina,  thus  being 
separated  from  lateral  area nnnulatus  Morley  (p.  121) 

-  Posterior  transverse  carina  of  propodeum  absent  centrally  (Fig.  29);  antenna  with  apical 

segments  black  but  with  a  subapical  white  band;  spiracular  area  confluent  with  lateral  area 

andrewi  sp.  n.  (p.  121) 

7  Males  and  females  with  tergites  2-5  with  yellow  band  along  hind  margin 8 

-  Females  with  tergites  2-5  with  paired  yellow  spots  or  the  males  either  with  paired  yellow  spots 

or  without  yellow  marks  at  all 9 

8  Metapleuron  centrally  with  conspicuous  longitudinal  wrinkles;  propodeum  with  a  tubercle 

below  spiracle  (Fig.  32) ;  flagellum  distally  uniformly  black;  anterolateral  part  of  mesoscutum 
without  yellow  marks geniculatus  Morley  (p.  124) 

-  Metapleuron  centrally  smooth;  propodeum  without  a  tubercle  below  spiracle  (Fig.  33); 

flagellum  distally  white-marked;  anterolateral  part  of  mesoscutum  yellow-marked 

nitidulus  Morley  (p.  131) 

9  Subalar  prominence  bearing  a  long  slender  back-curved  spine  (Fig.  34);  female  with  tergite  8 

posteriorly  extended  into  a  narrow  truncate  projection  (Fig.  35) talus  sp.  n.  (in  part)  (p.  135) 

-  Subalar  prominence  without  a  spine;  female  with  tergite  8  bluntly  rounded  apically  (Figs  36, 

37) 10 

10  Labium  with  glossae  very  long  and  slender,  extending  ventrally  a  distance  of  approximately  the 
height  of  the  eye;  occipital  carina  represented  by  a  short  vestige  at  its  junction  with 
hypostomal  carina  (Fig.  38);  propodeum  (at  least  part  behind  anterior  transverse  carina) 
yellow monticola  Morley(p.  130) 


120  I.  D.  GAULD  &  G.  A.  HOLLO  WAY 

Labium  with  glossae  short,  barely  projecting  below  head;  occipital  carina  present  ventrally,  at 
least  as  long  as  abscissa  of  hypostomal  carina  between  mandible  and  junction  with  occipital 
carina  (Fig.  39);  propodeum  not  or  only  partially  yellow-marked  behind  anterior  transverse 
carina 11 

1 1  Propodeum  with  anterior  and  posterior  transverse  carinae  strong ,  complete  and  almost  parallel 

to  each  other  (Fig.  40);  lateromedian  carinae  not  present  between  transverse  carina;  hind 

tibia  uniformly  reddish;  females  with  malar  space  brown sisyphus  sp.  n.  (p.  134) 

Propodeum  with  anterior  or  posterior  transverse  carina  weak  or  missing  in  part,  the  carinae  not 
parallel  and  often  with  discernible  traces  of  lateromedian  longitudinal  carinae  between  them 
(Fig .  41 ) ;  hind  tibia  proximally  pale-marked ;  females  with  malar  space  whitish 12 

12  Sternite  1  long,  extending  behind  spiracles  and  being  about  length  of  hind  coxa;  submetapleu- 

ral  carina  anteriorly  broadened  to  form  a  rectangular  flange  (Fig.  42);  mesoscutum  with  pair 

of  pale  stripes  extending  back  from  anterior  margin avitus  sp.  n.  (p.  123) 

Sternite  1  short,  reaching  at  most  (in  males)  to  level  of  spiracles,  and  being  distinctly  snorter 
than  length  of  hind  coxa;  submetapleural  carina  anteriorly  broadened  to  form  a  rounded 
lobe  (Fig.  43) ;  mesoscutum  with  a  median  pale  rectangular  mark cestus  sp.  n.  (p.  123) 

13  Hind  wing  with  distal  abscissa  of  Cu\  absent;  hind  tibia  with  posterior  margin  bearing  one  or 

more  spine-like  bristles 14 

Hind  wing  with  distal  abscissa  of  Cu\  present  in  membrane,  not  joining  nervellus;  hind  tibia 
with  posterior  margin  devoid  of  spine-like  bristles 22 

14  Tergite  2  of  gaster  entirely  white;  alitrunk  anteriorly  reddish  brown,  propodeum  infuscate, 

gaster  in  greater  part  black  (Fig.  58);  wings  uniformly  infumate;  antenna  of  female  black; 

female  with  posterior  margin  of  tergite  7  only  slightly  indented farrugiai  sp.  n.  (p.  126) 

Tergite  2  of  gaster  yellowish  brown,  or  dark  brown  or  pale  spotted  but  never  entirely  white 
(Figs  59-62) ;  alitrunk  variously  coloured,  if  brownish  then  concolorous  with  much  of  gaster ; 
wings  hyaline  or  apically  infumate;  antenna  of  female  with  white  bands;  female  with 
posterior  margin  deeply  indented  medially 15 

15  First  segment  of  gaster  exceptionally  long  and  slender,  the  Sternite  far  longer  than  length  of 

hind  coxa;  propodeum  with  first  and  second  lateral  areae  clearly  defined  and  separated  (Fig. 

44);  apex  of  fore  wing  of  female  narrowly  infumate celeus  sp.  n.  (p.  125) 

-  First  segment  of  gaster  stouter ,  the  Sternite  not  longer  than  hind  coxa ;  propodeum  with  first  and 

second  lateral  areae  confluent,  often  not  defined  laterally  (Fig.  45);  apex  of  fore  wing  not 
infumate 16 

16  Gaster  more  or  less  uniformly  yellowish  or  orange;  female  with  at  least  distal  two  flagellar 

segments  entirely  black  so  white  ruptive  mark  is  a  subapical  band 17 

-  Gaster  predominantly  black  with  white  maculae;  female  with  distal  end  of  flagellum  white 

except  for  tip  of  apical  segment  which  is  black 18 

17  Pronotum  convexly  produced  before  upper  hind  corner,  the  convexity  almost  pyramidal  in 

dorsal  view  (Fig.  46);  female  with  tergite  9  in  dorsal  view  transverse;  female  hind  tarsus 

black;  male  with  3r-m  present pineus  sp.  n.  (p.  132) 

Pronotum  weakly  convex  before  upper  hind  corner  (Fig.  47);  female  with  tergite  9  in  dorsal 
view  elongate;  female  hind  tarsus  yellow;  male  with  3r-m  absent. 

mogimbensis  Cheesman  (p.  129) 

18  Tergite  2-5  with  hind  margin  banded  with  yellow,  occasionally  with  bands  very  faint  so  tergite 

is  almost  unicolorous 19 

Tergite  2-5  with  yellow  spots  on  posterolateral  corners,  these  marks  not  confluent  centrally 21 

19  Propodeum  with  area  superomedia  distinct,  hexagonal,  with  only  posterolateral  sides  rather 

weak  (Fig.  48);  tergite  8  produced  into  long  slender  processes  (Fig.  50);  tergites  1-2  not 

clearly  whitish  or  yellow-marked species  A  (p.  137) 

Propodeum  with  area  superomedia  undefined  laterally  and  posteriorly  (Fig.  49);  tergite  8 
produced  into  blunt  or  moderately  long  processes  (Fig.  51);  tergites  1-2  clearly  whitish  or 
yellow-banded  posteriorly  (Fig.  59) 20 

20  Metapleuron  closely  punctate;  lower  face  distinctly  transverse  (Fig.  52);  hind  coxa  dorsally 

brown;  propodeum  in  lateral  aspect  brownish leeuwinensis  Turner  (p.  128) 

Metapleuron  virtually  smooth;  lower  face  longer  than  broad  (Fig.  53);  hind  coxa  entirely  black; 
propodeum  in  lateral  aspect  predominantly  whitish zebrus  sp.  n.  (p.  136) 

21  Pronotum  convexly  produced  before  upper  hind  corner;  propodeum  in  lateral  view  extensively 

whitish;  metapleuron  virtually  impunctate;  hind  tibia  with  an  indistinct  yellowish  mark 
proximally hinnuleus  Krieger  (p.  126) 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  121 

Pronotum  very  weakly  convex  before  upper  hind  corner;  propodeum  in  lateral  view  brown 
(Fig.  60);  metapleuron  closely  punctate;  hind  tibia  with  proximal  0-5  whitish 

toolangi  sp.  n.(p.  136) 

22  Subalar  prominence  bearing  a  long  slender  back-curved  spine  (Fig.  34);  female  flagellum  with  a 

subapical  white  band,  the  distal  5  or  so  segments  black talussp.  n.  (in  part)  (p.  135) 

Subalar  prominence  simple;  flagellum  with  apical  segments  white,  only  extreme  distal  apex  of 
last  segment  blackish  or  entirely  black 23 

23  Tergites  4  and  5  of  gaster  with  anterolateral  corners  broadly  white  (Fig.  62) ;  hind  coxa  in  profile 

rather  short  and  stout,  the  ovipositor  guide  reaching  to  about  the  centre  (Fig.  54);  process  on 

tergite  8  very  slender;  ovipositor  about  4  times  as  long  as  hind  tibia paluma  sp.  n.  (p.  132) 

-  Tergites  4  and  5  of  gaster  with  only  small  white  triangular  marks  in  posterolateral  corners  (Fig. 
61);  hind  coxa  in  profile  slender,  the  ovipositor  guide  reaching  about  0-3  of  its  length  (Fig. 
55);  process  on  tergite  8  moderately  slender;  ovipositor  about  5-5  times  as  long  as  hind  tibia 

ixion  sp.  n.(p.  128) 

Certonotus  andrewisp.  n. 

(Fig.  29) 

Female.  Medium  to  large-sized  species,  fore  wing  length  7-14  mm.  Labium  with  glossae  unspecialized. 
Lower  face  at  narrowest  point  1-0-1-1  times  as  broad  as  high;  malar  space  0-7-0-8  times  as  long  as  basal 
mandibular  width.  Occipital  carina  ventrally  more  than  twice  as  long  as  abscissa  of  hypostomal  carina 
between  it  and  base  of  mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  almost  flat,  in 
dorsal  view  barely  projecting  beyond  scutal  margin;  subalar  prominence  moderately  convex.  Scutellum 
transversely  striate,  crest  strong;  metapleuron  smooth  and  polished;  submetapleural  carina  anteriorly 
expanded  into  a  moderately  broad,  rather  long  lobe;  metanotum  with  very  strong  tooth  opposite  anterior 
end  of  lateral  carina.  Propodeum  moderately  short  with  anterior  transverse  carina  incomplete  centrally; 
posterior  transverse  carina  absent  centrally;  lateromedian  longitudinal  carina  present  only  before  anterior 
carina;  pleural  carina  complete,  but  weak  posteriorly;  area  superomedia  undefined;  area  spiracularis 
confluent  with  first  lateral  area;  first  and  second  lateral  area  separated  by  a  carina.  Fore  wing  with  3r-m 
converging  towards  2r-m,  joining  Rs  separately;  2m-cu  joining  M  0-25  to  0-50  from  3r-m  towards  2r-m. 
Hind  wing  with  distal  abscissa  of  Cu^  distinct  to  wing  margin.  Hind  tibia  with  posterior  margin  without 
spine-like  bristles.  Gaster  with  segment  1  short,  the  sternite  0-5-0-6  times  as  long  as  hind  coxa,  reaching  to 
level  of  spiracle.  Tergite  7  mediodorsally  without  any  indentation;  tergite  8  posteriorly  with  process  short 
and  rounded;  tergite  9  in  dorsal  view  transverse.  Ovipositor  projecting  beyond  apex  of  gaster  by  3-5  to  4-0 
times  length  of  hind  tibia. 

Coloration.  Orange-yellow  species  with  frons  centrally,  vertex,  mesoscutal  stripes,  most  of  gaster  and 
hind  legs  more  brownish  or  orange;  flagellum  black  with  subapical  white  band.  Pterostigma  brown,  wings 
hyaline. 

Male.  Similar  to  female. 

i 

REMARKS.  Very  like  C.  annulatus  which  it  resembles  in  colour,  venation,  possession  of  short  malar  space 
and  an  almost  flat  pronotum.  C.  andrewi  appears  to  be  a  more  southern  species  than  annulatus  as  it  has 
been  collected  in  southern  Queensland,  New  South  Wales  and  Victoria. 

MATERIAL  EXAMINED 

Holotype  $,  New  South  Wales:  Heathcote,  near  Sydney,  x.1979  (Holloway)  (AM). 

Paratypes.  Queensland:  1  <j>,  Brisbane,  xii.1972  (SedlaceK)  (TC);  4  <j>,  Mt  Glorious,  near  Brisbane, 
xii.1976  (Boucek)  (BMNH);  1  $,  6  cf ,  Mt  Glorious,  xi  (TC).  New  South  Wales:  1  $,  Cabbage  Tree  Ck, 
Clyde  Mtn,  ix.1979  (Naumann  &  Cardale)  (ANIC);  1  $,  Iluka,  Clarence  R.,  rain  forest,  xi.1970 
(McAlpine)  (AM);  1  $,  Warren,  ix.1982  (Holloway)  (AM).  Victoria:  1  $,  'Victoria'  (NMV). 

Certonotus  annulatus  Morley 
(Figs  28,  31) 

Certonotus  annulatus  Morley,  1913: 31;  Turner,  1919: 551.  LECTOTYPE  $ ,  QUEENSLAND  (BMNH),  here 
designated  [examined]. 

Female.  Moderately  large  species,  fore  wing  length  9-13  mm.  Labium  with  glossae  very  slightly  leng- 
thened. Lower  face  at  narrowest  point  0-9  times  as  broad  as  high;  malar  space  0-7-0-8  times  as  long  as  basal 
mandibular  width.  Occipital  carina  ventrally  about  as  long  as  abscissa  of  hypostomal  carina  between  it  and 


122  I.  D.  GAULD  &  G.  A.  HOLLOWAY 

mandibular  base.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  very  weakly  convex,  in  dorsal 
view  projecting  only  slightly  beyond  scutal  margin;  subalar  prominence  weakly  convex.  Scutellum 
punctate,  transverse  crest  distinct;  metapleuron  virtually  smooth  except  for  a  few  striae  ventrally  and 
posteriorly;  submetapleural  carina  anteriorly  expanded  into  a  broad  rounded  lobe  that  usually  bears 
concentric  striae;  metanotum  with  a  blunt  tooth  opposite  anterior  end  of  lateral  carina.  Propodeum 
moderately  long  with  anterior  transverse  carina  centrally  absent;  posterior  transverse  carina  centrally 
present;  lateromedian  longitudinal  carina  complete  only  anteriorly  and  with  stub  behind  anterior  trans- 
verse carina;  pleural  carina  complete;  area  superomedia  indicated,  but  incomplete  anteriorly  and  laterally; 
area  spiracularis  complete;  first  and  second  lateral  areae  separated,  distinctly  delineated.  Fore  wing  with 
3r-m  converging  towards  2r-m,  joining  Rs  separately;  2m-cu  joining  M  0-2-0-3  from  3r-m  towards  2r-m. 
Hind  wing  with  distal  abscissa  of  Cu\  distinct  to  wing  margin.  Hind  tibia  with  posterior  margin  without 
spine-like  bristles.  Gaster  with  segment  1  short,  the  sternite  0-5-0-6  times  as  long  as  hind  coxa,  reaching 
behind  level  of  spiracle.  Tergite  7  mediodorsally  with  a  short,  narrow  slit  on  posterior  margin;  tergite  8 
posteriorly  with  process  long  and  truncate;  tergite  9  in  dorsal  view  elongate  and  weakly  rounded 
posteriorly.  Ovipositor  projecting  beyond  apex  of  gaster  by  4-5  times  length  of  hind  tibia. 

Coloration.  Orange  species  with  face  and  orbits,  notauli,  subalar  prominences,  scutellar  margins  and 
hind  margins  of  gastral  tergites  more  yellowish;  flagellum  black  with  distal  10  or  so  segments  white. 
Pterostigma  blackish,  wings  very  weakly  infumate. 

Male.  Similar  to  female  but  with  fore  wing  length  8  mm;  malar  space  0-4  times  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  moderately  slender,  the  sternite  0-5  times  as  long  as  hind  coxa.  Apex  of 
gonosquama  flattened,  bearing  a  tuft  of  long  hairs.  Coloration  as  for  female. 

REMARKS.  C.  annulatus  is  rather  similar  to  and  probably  the  sister  species  of  C.  andrewi.  The  two  species 
are  fairly  easily  separated  by  the  characters  given  in  the  key. 
C.  annulatus  seems  to  be  a  north  Queensland  species. 

MATERIAL  EXAMINED 

Lectotype  $ ,  Queensland:  Kuranda  near  Cairns,  xii.1901  (BMNH). 

Queensland:  2  $  (paralectotypes),  Kuranda,  xii.1901,  iv.1902  (BMNH);  1  $,  Kuranda,  ii.1935  (Burns) 
(NMV);  1  £,  Mission  Beach,  near  Tully,  iv.1971  (Moulds}  (AM);  1  $,  2  cf ,  Moses  Ck,  4  km  N.  by  E.  Mt 
Finnigan  (15-47S  145-17E),  at  light,  x.1980  (Cardale)  (ANIC);  2  cf ,  Paluma  (19-OOS  146-12E),  900  m, 
Malaise  trap,  x.1980  (Frith)  (ANIC);  1  $,  Shipton's  Flat  (15-47S  145-14E),  at  light,  x.1980  (Cardale) 
(ANIC). 

Certonotus  apicalis  Morley 
(Fig.  23) 

Certonotus  apicalis  Morley,  1913:  31;  Turner,  1919:  551.  LECTOTYPE  $,  QUEENSLAND  (BMNH),  here 
designated  [examined] 

Female.  Large  species,  fore  wing  length  12-15  mm.  Labium  with  glossae  slightly  lengthened.  Lower  face  at 
narrowest  point  1-0-1-1  times  as  broad  as  high;  malar  space  0-7-0-8  times  as  long  as  basal  mandibular 
width.  Occipital  carina  ventrally  sinuous,  more  than  twice  as  long  as  abscissa  of  hypostomal  carina  between 
it  and  base  of  mandibles.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  convex,  in  dorsal  view 
projecting  as  a  rounded  protuberance;  subalar  prominence  quite  strongly  convex.  Scutellum  punctate, 
transverse  crest  weak;  metapleuron  with  fine  sparse  punctures;  submetapleural  carina  anteriorly  expanded 
into  a  broad  rounded  lobe;  metanotum  with  a  weak  swelling  in  front  of  lateral  carina.  Propodeum 
moderately  short  with  anterior  transverse  carina  complete;  posterior  transverse  carina  present  centrally; 
lateromedian  longitudinal  carina  present  before  anterior  carina  only;  pleural  carina  strong  anteriorly, 
weak  posteriorly;  area  superomedia  not  defined  laterally;  area  spiracularis  complete;  first  and  second 
lateral  areae  more  or  less  completely  separated,  the  latter  usually  undefined  externally.  Fore  wing  with 
3r-m  fused  anteriorly  with  2r-ra,  so  areolet  is  petiolate;  2m-cu  joining  M  0-20-0-25  basad  of  3r-m.  Hind 
wing  with  distal  abscissa  of  Cu\  distinct  to  wing  margin.  Hind  tibia  with  posterior  margin  without  spine-like 
bristles.  Gaster  with  segment  1  short,  the  sternite  0-5  times  as  long  as  hind  coxa,  not  reaching  level  of 
spiracle.  Tergite  7  mediodorsally  divided  about  0-5  times  its  length;  tergite  8  posteriorly  with  long  narrow 
process;  tergite  9  in  dorsal  view  transverse,  rounded  apically.  Ovipositor  projecting  beyond  apex  of  gaster 
by  7-5  or  more  times  length  of  hind  tibia. 

Coloration.  Yellowish  brown  species,  antenna  black  with  distal  segments  white,  the  apical  one  or  two 
segments  slightly  infuscate.  Pterostigma  black,  wings  weakly  infumate. 

Male.  Similar  to  female  but  with  fore  wing  length  9  mm;  malar  space  0-5  times  basal  mandibular  width; 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  123 

3r-m  fused  anteriorly  with  2r-m;  gaster  with  segment  1  short,  the  sternite  0-7  times  as  long  as  hind  coxa, 
apex  of  gonosquama  flattened,  with  a  tuft  of  long  close  hairs.  Colour  similar  to  female  but  pterostigma  dark 
brown. 

REMARKS.  The  females  of  this  species  are  easily  distinguished  by  their  very  long  ovipositors,  the  apices  of 
which  are  fairly  bluntly  pointed.  The  fore  femur  is  also  rather  distinctive,  being  slender  proximally,  then 
abruptly  inflated  proximocentrally.  This  is  most  similar  to  C.  humeralifer  and  the  two  are  probably 
sister-taxa. 

C.  apicalis  is  a  tropical  species,  only  recorded  from  Queensland. 

MATERIAL  EXAMINED 

Lectotype  $ ,  Queensway:  Kuranda  near  Cairns,  iv.1902  (BMNH). 

Queensland:  3  $  (paralectotypes),  same  data  as  holotype  (BMNH);  1  d",  Kuranda,  v-vi.1913  (Turner) 
(BMNH);  1  $,  Paluma,  mv  lamp,  i.1970  (Holloway)  (AM). 

Certonotus  avitus  sp.  n. 

(Figs  41,  42) 

Female.  Medium-sized  species,  fore  wing  length  8-10  mm.  Labium  with  glossae  unspecialized.  Lower  face 
at  narrowest  point  1-0  times  as  broad  as  high;  malar  space  0-4-0-6  times  as  long  as  basal  mandibular  width. 
Occipital  carina  ventrally  sinuous,  more  than  3  times  as  long  as  abscissa  of  hypostomal  carina  between  it 
and  base  of  mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  almost  flat,  in  dorsal  view 
barely  projecting  beyond  scutal  margin;  subalar  prominence  weakly  convex.  Scutellum  finely  punctate, 
crest  strong;  metapleuron  finely  punctate,  obsolescent  striae  posteriorly;  submetapleural  carina  anteriorly 
expanded  into  a  rectangular  elongate  flange;  metanotum  laterally  produced  into  a  stout  tooth  in  front  of 
lateral  carina.  Propodeum  moderately  long  with  anterior  transverse  carina  incomplete  centrally;  posterior 
transverse  carina  absent  except  laterally;  lateromedian  longitudinal  carina  present  only  before  anterior 
carina;  pleural  carina  incomplete  posteriorly;  area  superomedia  indistinct;  area  spiracularis  complete;  first 
and  second  lateral  areae  separated,  the  second  not  defined  laterally.  Fore  wing  with  3r-m  converging 
towards  2r-m,  widely  separated  on  Rs;  2m-cu  joining  M  0-3  from  3r-m  towards  2r-m.  Hind  wing  with  distal 
abscissa  of  Cui  distinct  to  wing  margin.  Hind  tibia  with  posterior  margin  without  spine-like  bristles.  Gaster 
with  segment  1  long,  the  sternite  1-2  times  as  long  as  hind  coxa,  reaching  well  behind  level  of  spiracle. 
Tergite  7  mediodorsally  with  indentation  to  0-25  of  tergite  length;  tergite  8  posteriorly  with  short,  rounded 
process;  tergite  9  in  dorsal  view  transverse.  Ovipositor  projecting  beyond  apex  of  gaster  by  5  times  length 
of  hind  tibia. 

Coloration.  Flagellum  black,  distal  apex  white;  head  brown,  orbits  entirely  and  all  of  lower  face  whitish; 
alitrunk  dark  brown,  notaular  stripes,  subalar  prominence,  tegula,  scutellum,  postscutellum,  mesopleural 
stripe  and  posterior  part  of  propodeum/metapleuron  yellow.  Gaster  reddish  brown,  tergites  laterally 
yellow  and  with  posterolateral  yellow  spots.  Tergite  1  anteriorly  yellowish.  Anterior  two  pairs  of  coxae 
whitish,  hind  ones  brown  with  white  apices;  all  tibiae  and  tarsi  infuscate;  femora  brown,  distally  pale. 
Pterostigma  dark  brown;  wings  hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  8  mm;  malar  space  0-3-0-4  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  slender,  the  sternite  1-0  times  as  long  as  hind  coxa.  Gonosquama  with 
apex  slightly  flattened,  with  a  tuft  of  close  moderately  long  hairs.  Similar  in  colour  to  female. 

REMARKS.  C.  avitus  is  a  distinctive  species  with  a  very  characteristic  submetapleural  carina.  For  its  size  this 
species  has  a  rather  large  areolet  and  a  fairly  long  first  sternite. 
It  is  known  from  eastern  Australia,  from  southern  Queensland  to  Victoria. 

MATERIAL  EXAMINED 

Holotype  $ ,  New  South  Wales:  Mt  Tomah,  Blue  Mtns,  viii.1979  (Rodd)  (AM). 

Paratypes.  Queensland:  1  $ ,  Eungella,  xi  (TC);  1  $ ,  Mt  Glorious,  i  (TC);  1  cf ,  Mt  Nebo,  viii  (TC);  1  $ , 
Mt  Tamborine,  ix-x.1978  (Galloway)  (BMNH).  Victoria:  1  cT,  Healesville,  xi.1943  (NMV);  1  $,  King 
Lake,  x.1953  (Burns).  (NMV);  1  C?,  King  Lake,  x.1954  (Burns)  (NMV);  4  $>,  1  cf,  Toolangi,  xi.1982 
(Farrugia)  (BMNH). 

Certonotus  cestus  sp.  n. 

(Figs  37,  43) 

Female.  Medium-sized  species,  fore  wing  length  9  mm.  Labium  with  glossae  unspecialized.  Lower  face  at 
narrowest  point  1-1  times  as  broad  as  high;  malar  space  1-0  times  as  long  as  basal  mandibular  width. 


124  I.  D.  GAULD  &  G.  A.  HOLLO  WAY 

Occipital  carina  ventrally  longer  than  abscissa  of  hypostomal  carina  between  it  and  lobe  of  mandible. 
Upper  part  of  pronotum,  slightly  before  posterior  cornet,  weakly  convex,  in  dorsal  view  barely  protruding 
beyond  scutal  margin;  subalar  prominence  quite  strongly  convex.  Scutellum  coarsely  punctate,  crest 
strong;  metapleuron  small  and  polished;  submetapleural  carina  anteriorly  expanded  into  a  rounded  lobe, 
metanotum  barely  produced  before  lateral  carina.  Propodeum  moderately  long  with  anterior  transverse 
carina  obsolescent  centrally;  posterior  transverse  carina  weak  but  more  or  less  complete;  lateromedian 
longitudinal  carina  present  anteriorly  and  weakly  between  transverse  carina;  pleural  carina  anteriorly 
strong,  posteriorly  weaker  or  absent;  area  superomedia  more  or  less  distinct,  not  defined  anteriorly;  area 
spiracularis  complete;  first  and  second  lateral  areae  separated,  the  first  very  short,  the  second  ill-defined 
externally.  Fore  wing  with  3r-m  converging  towards  2r-ra,  joining  Rs  separately;  2m-cu  joining  M  opposite 
3r-m.  Hind  wing  with  distal  abscissa  of  Cu\  distinct  to  wing  margin.  Hind  tibia  with  posterior  margin  with 
two  spine-like  bristles.  Gaster  with  segment  1  short  and  broad  posteriorly,  the  sternite  0-5  times  as  long  as 
hind  coxa,  reaching  almost  to  level  of  spiracle.  Tergite  7  mediodorsally  with  broad  indentation;  tergite  8 
posteriorly  short  and  rounded;  tergite  9  in  dorsal  view  very  broad.  Ovipositor  projecting  beyond  apex  of 
gaster  by  6  times  length  of  hind  tibia. 

Coloration.  Very  dark  brown  species,  flagellum  with  a  white  subapical  band;  facial,  frontal  and  genal 
orbits,  upper  and  lower  margin  of  pronotum,  mesoscutum  in  a  central  quadrate  spot  and  indistinctly 
laterally,  scutellum,  postscutellum,  subalar  prominence,  anterior  and  posterior  mesopleural  spots,  most  of 
hind  part  of  propodeum  and  metapleuron  whitish.  Gaster  dark  brown  with  large  anterolateral  spots. 
Anterior  coxa  whitish,  hind  coxa  brown  with  whitish  mark.  Pterostigma  blackish,  wings  hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  9  mm;  malar  space  0-7  times  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  quite  stout,  the  sternite  0-5  times  as  long  as  hind  coxa.  Apex  of 
gonosquama  with  a  small  lobe  that  bears  scattered  long  hairs.  Colour  similar  to  female  but  face  entirely 
pale. 

REMARKS.  This  species  is  readily  recognizable  by  its  colour  pattern.  Structurally  it  seems  to  be  related  to  the 
C.  humeralifer-group  as  it  has  tergite  7  very  deeply  divided.  Tergite  8  is  less  pronounced  apically  in  this 
species  than  others  in  the  humeralifer-group. 
It  is  known  from  Queensland  and  New  South  Wales. 

HOST  RECORD.  Buprestidae:  Diadoxussp.  (ANIC). 

MATERIAL  EXAMINED 

Holotype  $ ,  New  South  Wales:  State  Forest  854,  viii.1952  (Martin)  (ANIC),  parasite  of  Diadoxus. 
Paratype.  Queensland:  1  cf ,  viii.1926  (Jarvis)  (BMNH). 

Certonotus  geniculatus  Morley 
(Fig.  32) 

Certonotus  geniculatus  Morley,  1913:  28.  LECTOTYPE  $,  VICTORIA  (BMNH),  here  designated  [ex- 
amined] . 

Female.  Large  species,  fore  wing  length  12-17  mm.  Labium  with  glossae  slightly  lengthened.  Lower  face  at 
narrowest  point  1-1-1-2  times  as  broad  as  high;  malar  space  0-9-1-0  times  as  long  as  basal  mandibular 
width.  Occipital  carina  ventrally  about  as  long  as  abscissa  of  hypostomal  carina  between  it  and  base  of 
mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  very  weakly  convex,  in  dorsal  view 
barely  projecting  beyond  scutal  margin;  subalar  prominence  weakly  convex.  Scutellum  punctate  with 
strong  transverse  crest;  metapleuron  longitudinally  striate;  submetapleural  carina  anteriorly  expanded 
into  a  moderately  broad  lobe;  metanotum  produced  into  a  tooth  opposite  anterior  end  of  lateral  carina. 
Propodeum  quite  short  with  anterior  transverse  carina  incomplete  centrally;  posterior  transverse  carina 
vestigial;  lateromedian  longitudinal  carina  present  only  as  vestige  anteriorly;  pleural  carina  strongly  raised 
into  tubercle  below  spiracle,  posteriorly  obsolescent;  area  superomedia  undefined;  area  spiracularis  not 
completely  delineated  posteriorly;  first  and  second  lateral  areae  separated,  but  not  clearly  delineated 
internally  or  laterally.  Fore  wing  with  3r-m  and  2r-m  converging  but  not  joining  anteriorly.  Hind  wing  with 
distal  abscissa  of  Cu\  distinct  to  hind  margin.  Hind  tibia  with  posterior  margin  without  spine-like  bristles. 
Gaster  with  segment  1  long  and  narrow,  the  sternite  1-0-1-3  times  as  long  as  hind  coxa,  reaching  behind 
level  of  spiracle.  Tergite  7  mediodorsally  with  very  small  indentation  on  posterior  margin;  tergite  8 
posteriorly  short  and  rounded.  Tergite  9  in  dorsal  view  broad.  Ovipositor  projecting  beyond  apex  of  gaster 
by  5-5  to  7-0  times  length  of  hind  tibia. 

Coloration.  Antenna  black;  head  black,  lower  face,  frontal  and  genal  orbits  yellow.  Alitrunk  red-brown 
with  periphery  of  sclerites  infuscate,  only  tegula,  scutellum  posteriorly,  postscutellum  and  hind  end  of 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  125 

propodeum  yellow.  Gaster  reddish  brown,  tergite  1  anteriorly,  laterally  and  posteriorly  yellow,  other 
tergites  with  lateral  and  posterior  margins  yellow.  Fore  leg  yellow,  proximal  part  of  femur  black;  mid  leg 
yellow,  coxa  basally,  femur  proximally  and  tarsus  black.  Hind  leg  black,  coxa  distally,  trochanter 
segments,  distal  apex  of  femur,  base  and  apex  of  tibia  yellow.  Pterostigma  dark  brown,  wings  virtually 
hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  11  mm;  malar  space  0-7-0-9  times  basal  mandibular 
width;  3r-m  present;  gaster  with  segment  1  very  slender,  the  sternite  1-3  times  as  long  as  hind  coxa.  Apex  of 
gonosquama  flattened  with  a  dense  tuft  of  long  hairs.  Colour  similar  to  female. 

REMARKS.  C.  geniculatus  belongs  to  the  rufescens-subgroup  of  the  tasmaniensis-group.  It  is  probably  the 
sister-species  of  C.  rufescens.  Both  have  the  prementum  and  glossae  somewhat  lengthened,  have  a  very  flat 
face,  the  anterior  portion  of  the  lateral  carina  strongly  raised  and  the  pleural  carina  raised  into  a  tubercle 
below  the  propodeal  spiracle.  C.  geniculatus  is  most  easily  distinguished  by  the  possession  of  a  longitudin- 
ally striate  metapleuron. 
This  is  a  southern  species,  only  recorded  from  Victoria. 

MATERIAL  EXAMINED 

Lectotype  $,  Victoria:  Nulla  Wurren,  near  Berwick  (BMNH). 

Victoria:  1  $,  2  O"  (paralectotypes),  same  data  as  lectotype  (BMNH);  1  9>  Buckland  River,  xi.1964 
(Neboiss)  (NMV);  1  cf ,  Fernshaw  (NMV);  1  $,  1  cf ,  Trafalgar  (NMV). 

Certonotus  celeussp.  n. 

(Fig.  44) 

Female.  Medium-sized  species,  fore  wing  length  8-9  mm.  Labium  with  glossae  slightly  lengthened.  Lower 
face  at  narrowest  point  0-9  times  as  broad  as  high;  malar  space  1-3-1-4  times  as  long  as  basal  mandibular 
width.  Occipital  carina  ventrally  slightly  longer  than  abscissa  of  hypostomal  carina  between  it  and  base  of 
mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  very  weakly  convex,  in  dorsal  view 
projecting  slightly  beyond  mesoscutal  margin;  subalar  prominence  moderately  convex.  Scutellum  sparsely 
punctate  with  strong  crest  posteriorly;  metapleuron  smooth,  very  sparsely  punctate;  submetapleural  carina 
anteriorly  strongly  expanded  into  a  quadrate  lobe  that  is  abruptly  narrowed  posteriorly;  metanotum  with  a 
weak  lateral  tooth.  Propodeum  quite  long  with  anterior  transverse  carina  complete;  posterior  transverse 
carina  complete;  lateromedian  longitudinal  carina  present  only  anterior  to  anterior  carina;  pleural  carina 
complete;  area  superomedia  not  defined;  area  spiracularis  clearly  delineated;  first  and  second  lateral  areae 
distinct,  separated.  Fore  wing  with  3r-m  converging  towards  2r-m,  joining  Rs  separately;  2m-cu  joining  M 
0-3  from  3r-m  towards  2r-m.  Hind  wing  with  distal  abscissa  of  Cu\  absent.  Hind  tibia  with  posterior  margin 
with  one  spine-like  bristle.  Gaster  with  segment  1  very  long  and  narrow,  the  sternite  1-2-1-6  times  as  long 
as  hind  coxa,  reaching  well  behind  level  of  spiracle.  Tergite  7  mediodorsally  with  shallow  indentation; 
tergite  8  posteriorly  short  and  rounded;  tergite  9  in  dorsal  view  short  and  pointed.  Ovipositor  projecting 
beyond  apex  of  gaster  by  4-5  times  length  of  hind  tibia. 

Coloration.  Red-brown  species,  flagellum  black  with  subapical  white  band,  lower  face  yellowish, 
anterior  leg  orange.  Pterostigma  dark  brown;  wings  hyaline,  apex  of  fore  wing  strongly  infumate. 

Putative  male.  Similar  to  female  but  with  fore  wing  length  8  mm;  malar  space  0-6  times  basal  mandibular 
width;  3r-m  present;  propodeal  carinae  weaker;  gaster  with  segment  1  slender,  the  sternite  0-9  times  as 
long  as  hind  coxa.  Apex  of  gonosquama  rounded,  with  scattered  fine  hairs.  Pale  orange,  antenna  black, 
head  yellowish  and  wings  uniformly  hyaline. 

REMARKS.  A  distinctive  species  on  account  of  the  infumate  tip  to  the  fore  wing  of  the  female,  the  very  long 
petiole,  long  hind  coxa  and  rather  flat  face.  It  is  probably  related  to  C.  rufescens  though  it  is  the  only  species 
in  this  subgroup  without  the  distal  abscissa  of  Cu\.  The  male,  here  tentatively  associated,  has  similarly  long 
coxae  and  a  rather  flat  face  but  has  no  clear  posterior  transverse  propodeal  carina  nor  has  the  wing  apices 
infumate. 
It  is  only  known  from  Queensland. 

MATERIAL  EXAMINED 

Holotype  $,  Queensland:  Baldy  Mtn  Rd,  via  Atherton,  Malaise  trap,  vi.1981  (Brown)  (QM). 

Paratypes.  Queensland:  1  $,  same  data  as  holotype  (BMNH);  1  cf,  Eungella  Nat.  Park,  xi.1976 
(Boucek)  (BMNH);  1  $,  Windsor  Tableland,  iii.1981  (Storey)  (BMNH). 


126  I.  D.  GAULD  &  G.  A.  HOLLO  WAY 

Certonotusfarrugiaisp.  n. 

(Fig.  58) 

Female.  Medium-sized  species,  fore  wing  length  6-10  mm.  Labium  with  glossae  unspecialized.  Lower  face 
at  narrowest  point  1-1-1-2  times  as  broad  as  high;  malar  space  1-2-1-3  times  as  long  as  basal  mandibular 
width.  Occipital  carina  ventrally  as  long  as  abscissa  of  hypostomal  carina  between  it  and  base  of  mandible. 
Upper  part  of  pronotum,  slightly  before  posterior  corner,  almost  flat,  in  dorsal  view  barely  projecting 
beyond  mesoscutal  margin;  subalar  prominence  quite  strongly  convex  and  medially  raised  in  dorsal  aspect. 
Scutellum  smooth,  virtually  impunctate  with  weak  transverse  carina  apically;  metapleuron  anteriorly 
smooth,  posteriorly  with  few  coarse  punctures;  submetapleural  carina  anteriorly  expanded  into  a  broad 
rounded  lobe;  metanotum  with  a  small  lateral  tooth  before  vestige  of  lateral  carina.  Propodeum 
moderately  short  with  anterior  transverse  carina  complete;  posterior  transverse  carina  absent;  laterome- 
dian  longitudinal  carina  absent;  pleural  carina  present,  posteriorly  obsolescent;  area  superomedia 
undefined;  area  spiracularis  defined  except  medially;  first  and  second  lateral  areae  confluent.  Fore  wing 
with  3r-m  converging  towards  2r-m,  joining  latter  at  Rs  or  sometimes  slightly  separated;  2m-cu  joining  M 
slightly  basad  of  3r-m.  Hind  wing  with  distal  abscissa  of  Cu\  absent.  Hind  tibia  with  posterior  margin  with 
one  spine-like  bristle.  Caster  with  segment  1  short  and  broad,  the  sternite  0-4-0-6  times  as  long  as  hind 
coxa,  reaching  to  level  of  spiracle.  Tergite  7  mediodorsally  with  wide  indentation;  tergite  8  posteriorly 
short  and  broadly  rounded;  tergite  9  in  dorsal  view  short  and  broad.  Ovipositor  projecting  beyond  apex  of 
gaster  by  3-0-3-5  times  length  of  hind  tibia. 

Coloration.  Head  and  anterior  part  of  alitrunk  reddish  brown;  antenna,  much  of  metapleuron  and 
propodeum  black;  gaster  black,  tergite  2  entirely  and  posterior  margins  of  tergites  3+  white;  anteriorly 
red-brown,  partially  infuscate;  hind  leg  mainly  black.  Pterostigma  black,  wings  strongly  infumate. 

Male.  Similar  to  female  but  with  fore  wing  length  6  mm;  malar  space  0-8  times  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  stout,  the  sternite  0-6  times  as  long  as  hind  coxa.  Apex  of  gaster 
rounded,  bearing  long  scattered  hairs.  Similar  in  colour  to  female  but  with  gaster  entirely  red;  mid  leg 
brownish. 

REMARKS.  C.  farrugiai  is  probably  the  most  distinct  species  in  the  genus  on  account  of  its  striking  colour 
pattern  which  resembles  that  of  a  number  of  other  unrelated  species  of  Hymenoptera  occurring  in 
south-eastern  Australia.  Structurally  it  is  also  distinctive  in  having  the  posterior  margin  of  tergite  7  only 
weakly  indented. 
This  species  occurs  from  south-eastern  Queensland  to  Victoria. 

MATERIAL  EXAMINED 

Holotype  $,  Victoria:  Toolangi,  xii.1982  (Farrugia)  (AM). 

Paratypes.  Queensland:  4  $ ,  Mt  Glorious,  xi-iii  (TC);  1  $ ,  Mt  Nebo,  500  m,  iii  (TC);  1  $ ,  Tambourine, 
x.1977  (Galloway)  (BMNH);  1  tf ,  Mt  Tambourine,  iv.  1935  (Turner)  (BMNH);  1  $ ,  2  cT,  Mt  Tambourine, 
x-xii  (TC).  Victoria:  2  $,  same  data  as  holotype  (BMNH). 

Certonotus  hinnuleus  Krieger  comb.  rev. 

Certonotus  hinnuleus  Krieger,  1901:  123;  Turner,  1919:  551.  Holotype  $,  NEW  SOUTH  WALES  (MNHU) 

[not  examined] . 

Certonotus  n.  sp.;  Fullaway,  1942:  244. 
Asperellus  hinnuleus  (Krieger)  Townes  et  al. ,  1961: 114. 

Female.  Fairly  small  species,  fore  wing  length  4-7  mm.  Labium  with  glossae  unspecialized.  Lower  face  at 
narrowest  point  1-2  times  as  broad  as  high;  malar  space  1-2-1-4  times  as  long  as  basal  mandibular  width. 
Occipital  carina  ventrally  almost  twice  as  long  as  abscissa  of  hypostomal  carina  between  it  and  mandible 
base.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  very  convex,  in  dorsal  view  appearing  as  a 
conical  projection;  subalar  prominence  moderately  convex.  Scutellum  sparsely  punctate  with  strong 
transverse  keel;  metapleuron  smooth  and  polished;  submetapleural  carina  anteriorly  expanded  into  a 
broad  triangular  lobe;  metanotum  without  a  tooth.  Propodeum  short  with  anterior  transverse  carina 
complete,  close  to  anterior  margin;  posterior  transverse  carina  absent;  lateromedian  longitudinal  carina 
present  before  anterior  transverse  carina;  pleural  carina  present  anteriorly,  posteriorly  absent;  area 
superomedia  not  delineated;  area  spiracularis  not  delineated  internally;  first  and  second  lateral  areae 
indistinctly  delineated,  confluent.  Fore  wing  with  3r-m  converging  towards  2r-m,  sometimes  forming  a 
petiolate  areolet  or  joining  at  one  point  on  Rs;  2m-cu  joining  M  at  3r-m  or  slightly  basad.  Hind  wing  with 
distal  abscissa  of  Cui  absent.  Hind  tibia  with  posterior  margin  with  one  spine-like  bristle.  Gaster  with 
segment  1  short  and  broad,  the  sternite  0-6  times  as  long  as  hind  coxa,  reaching  to  level  of  spiracle.  Tergite  7 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  127 

mediodorsally  with  large  indentation;  tergite  8  posteriorly  short,  broad,  truncate;  tergite  9  in  dorsal  view 
short,  rounded  posteriorly.  Ovipositor  projecting  beyond  apex  of  gaster  by  5-0-5-5  times  length  of  hind 
tibia. 

Coloration.  Antenna  black,  distal  flagellar  segments  white;  head  white,  vertex,  interocellar  area  and 
frons  centrally  whitish;  mesoscutum  brownish  or  blackish  with  a  yellow  central  quadrate  mark;  most  of 
pronotum,  tegula,  subalar  prominence,  metapleuron  and  propodeum  almost  entirely  yellowish;  meta- 
pleuron  brownish.  Gaster  dark  brown  with  lateral  margins  and  spots  near  posterolateral  corners  yellow; 
legs  brownish;  tibia  proximally  and  femur  distally  somewhat  paler;  fore  coxa  whitish.  Pterostigma 
blackish,  wings  very  weakly  infumate. 

Male.  Similar  to  female  but  with  fore  wing  length  3-6  mm;  malar  space  0-9-1-0  times  basal  mandibular 
width;  3r-m  present;  gaster  with  segment  1  quite  stout,  the  sternite  0-5  times  as  long  as  hind  coxa;  apex  of 
gonosquama  quite  weakly  sclerotized,  slightly  flattened,  bearing  long  fine  hairs.  Colour  similar  to  female 
but  mid  coxa  yellow. 

REMARKS.  Certonotus  hinnuleus  belongs  to  the  leeuwinensis-group.  It  is  the  most  distinctive  taxon  in  the 
group  on  account  of  the  swollen  pronotal  corner.  Structurally  it  is  most  closely  related  to  C.  zebrus  from 
which  it  differs  in  having  a  longer  ovipositor  and  shorter  stouter  petiole.  In  Australia  it  has  only  been 
recorded  from  New  South  Wales  and  Victoria.  The  female  from  Western  Australia  referred  to  by  Morley 
(1913)  is  a  distinct  species,  C.  ixion.  Townes  et  al.  (1961)  note  that  this  species  occurs  in  New  Caledonia. 

MATERIAL  EXAMINED 

New  Caledonia:  2  $ ,  7  km  SE.  La  Foa,  i.  1945  (Remington)  (TC)  (compared  with  type);  1  d",  hills  behind 
Noumea,  x.1940  (Williams)  (TC).  Queensland:  1  C?,  N.  slope,  Bluff  Range,  Biggenden,  viii.1976  (Frauca) 
(ANIC);  1  C?,  Broken  R.,  near  Eungella,  xii.1961  (McAlpine  &  Lossiri)  (AM);  1  $,  1  cf,  Kuranda, 
v-vi.1913  (Turner)  (BMNH);  1  C?,  Mackay,  1909  (Turner)  (BMNH);  3  C?,  Mt  Cootha,  iv-v  (TC);  1  C?,  Mt 
Glorious,  i  (TC);  2  d" ,  Mt  Tambourine,  xi.1977  (Galloway)  (BMNH). 

Certonotus  humeralifer  Krieger 
(Figs  24,  26,  27,  30) 

Certonotus  humeralifer  Krieger,  1901:  121.  Lectotype  $,  NEW  SOUTH  WALES  (MNHU),  designated  by 
Townes  et  al.,  1961: 113  [examined]. 

Female.  Medium  to  large  species,  fore  wing  length  7-14  mm.  Labium  with  glossae  unspecialized.  Lower 
face  at  narrowest  point  1-2-1-3  times  as  broad  as  high;  malar  space  0-7-0-8  times  as  long  as  basal 
mandibular  width.  Occipital  carina  ventrally  sinuous,  at  least  twice  as  long  as  abscissa  of  hypostomal  carina 
between  it  and  base  of  mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  very  strongly 
swollen,  in  dorsal  view  pyramidal,  projecting;  subalar  prominence  strongly  convex.  Scutellum  punctate, 
crest  distinct,  metapleuron  smooth  and  polished;  submetapleural  carina  anteriorly  expanded  into  a  tooth 
before  lateral  carina.  Propodeum  moderately  short  with  anterior  transverse  carina  weak  centrally  but 
usually  complete;  posterior  transverse  carina  absent;  lateromedian  longitudinal  carina  present  anteriorly; 
pleural  carina  strong  anteriorly,  posteriorly  vestigial;  area  superomedia  not  delineated;  area  spiracularis 
not  clearly  delineated  posteriorly;  first  and  second  lateral  area  confluent,  weakly  defined  laterally.  Fore 
wing  with  3r-m  fused  anteriorly  with  2r-m,  so  areolet  petiolate;  2m-cu  joining  M-basad  by  0-2  from  3r-m. 
Hind  wing  with  distal  abscissa  of  Cui  distinct  to  wing  margin.  Hind  tibia  with  posterior  margin  with  one 
spine-like  bristle.  Gaster  with  segment  1  short,  the  sternite  0-5  times  as  long  as  hind  coxa,  reaching  to  level 
of  spiracle.  Tergite  7  mediodorsally  with  wide  indentation  narrowing  to  slit  to  0-5  of  tergite  length;  tergite  9 
in  dorsal  view  short,  rectangular.  Ovipositor  projecting  beyond  apex  of  gaster  by  5  times  length  of  hind 
tibia. 

Coloration.  Bright  yellow  species;  flagellum,  interocellar  area,  mesoscutum  in  central  spot  and 
scutoscutellar  groove  black;  hind  tarsus  infuscate.  Pterostigma  black,  wings  hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  7-8  mm;  malar  space  0-6  times  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  quite  stout,  the  sternite  0-5  times  as  long  as  hind  coxa;  apex  of 
gonosquama  strongly  flattened,  with  periphery  bearing  only  fine  sparse  hairs.  Similar  in  colour  to  female. 

REMARKS.  The  bright  yellow  ground  colour  of  this  species  and  the  conical  pronotal  process  distinguish  C. 
humeralifer  from  other  Australian  Certonotus  species.  C.  humeralifer  belongs  to  the  humeralifer-group;  it 
appears  to  be  most  closely  related  to  C.  apicalis.  Both  have  a  characteristically  specialized  fore  femur. 
C.  humeralifer  has  been  collected  in  Queensland  and  New  South  Wales. 


128  I.  D.  GAULD  &  G.  A.  HOLLO  WAY 

MATERIAL  EXAMINED 

Lectotype  $,  New  South  Wales  (MNHU). 

Queensland:  1  cf ,  Brisbane,  1956  (Ken)  (NMV);  2  $,  Fraser  Island,  ix.1930  (BMNH);  1  $,  1  cf ,  Mt 
Glorious,  650  m,  x-xiii  (TC);  1  $,  Mt  Nebo,  500  m,  iii  (TC);  5  $,  1  cf,  Mt  Tambourine,  x-xi.1977 
(Galloway)  (BMNH);  1  $,  Mt  Tambourine,  xii.1911  (Hacker)  (BMNH);  2  $,  Mt  Tambourine,  xi-xii 
(TC);  1  cf ,  Toowoomba,  iii  (TC).  New  South  Wales:  1  $,  Terrigal,  1900  (Froggatt)  (BMNH);  1  $,  'New 
South  Wales' (NMV). 

Certonotus  ixion  sp.  n. 

(Figs  55,  61) 

Female.  Medium-sized  species,  fore  wing  length  8  mm.  Labium  with  glossae  unspecialized.  Lower  face  at 
narrowest  point  1-2  times  as  broad  as  high;  malar  space  0-9  times  as  long  as  basal  mandibular  width. 
Occipital  carina  ventrally  slightly  longer  than  abscissa  of  hypostomal  carina  between  it  and  base  of 
mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  weakly  convex,  in  dorsal  view  parallel 
with  mesoscutal  margin;  subalar  prominence  moderately  convex.  Scutellum  closely,  finely  punctate  with 
strong  transverse  crest;  metapleuron  with  close  shallow  punctures;  submetapleural  carina  anteriorly 
abruptly  expanded  into  a  broad  lobe.  Propodeum  moderately  short  with  anterior  transverse  carina 
complete;  posterior  transverse  carina  almost  complete;  lateromedian  longitudinal  carina  absent  except  for 
vestiges  before  anterior  carina;  pleural  carina  obsolescent  posteriorly;  area  superomedia  not  delimited 
laterally;  area  spiracularis  almost  complete;  first  and  second  lateral  areae  barely  delineated  laterally, 
confluent.  Fore  wing  with  3r-m  strongly  converging  towards  2r-m,  joining  Rs  separately;  2m-cu  joining  M 
at  3r-m.  Hind  wing  with  distal  abscissa  of  Cu\  present  over  distal  0-5  to  wing  margin,  not  joined  to  first 
abscissa  of  Cu\.  Hind  tibia  with  posterior  margin  with  a  small  spine-like  bristle.  Gaster  with  segment  1  long 
and  narrow,  the  sternite  0-8  times  as  long  as  hind  coxa,  reaching  behind  level  of  spiracle.  Tergite  7 
mediodorsally  with  narrow  indentation  to  0-5  of  length;  tergite  8  posteriorly  short  and  truncate;  tergite  9  in 
dorsal  view  broad.  Ovipositor  projecting  beyond  apex  of  gaster  by  5-5  times  length  of  hind  tibia. 

Coloration.  Antenna  blackish,  apex  (except  extreme  distal  part  of  last  segment)  whitish;  head  reddish 
brown,  lower  face,  frontal  orbits  and  most  of  genae  whitish;  alitrunk  reddish,  peripherally  darker; 
scutellum  posteriorly,  postscutellum,  tegula,  subalar  prominence  and  upper  margin  of  pronotum  yellow- 
ish. Gaster  dark  red-brown;  tergites  1-6  with  posterolateral  triangular  yellow  marks,  tergites  2-3  with 
anterolateral  pale  spots.  Legs  predominantly  reddish  brown,  anterior  two  pairs  of  coxae  and  distal  apices 
of  fore  and  mid  femora  whitish.  Pterostigma  red-brown,  wings  hyaline. 

Male.  Quite  similar  to  female  but  with  fore  wing  length  5-7  mm;  malar  space  0-6  times  basal  mandibular 
width;  3r-m  present;  gaster  with  segment  1  slender,  the  sternite  1-0  times  as  long  as  hind  coxa;  apex  of 
gaster  with  a  small  lobe  bearing  long  scattered  hairs.  Colour  similar  to  female  but  with  mesopleuron 
white-marked  and  pale  maculae  on  gaster  smaller,  rather  inconspicuous,  flagellum  entirely  black. 

REMARKS.  The  species  is  one  of  the  three  Certonotus  that  have  incomplete  distal  abscissa  of  Cu\,  It  is  most 
similar  to  C.  paluma  from  which  it  differs  strikingly  in  colour  pattern,  length  of  ovipositor  and  shape  of 
tergite  8.  The  coxae  are  more  elongate  than  in  many  other  species  and  the  sculpture  of  the  alitrunk  is 
coarser  than  that  of  C.  paluma. 

C.  ixion  has  been  collected  in  Queensland  and  Victoria. 

MATERIAL  EXAMINED 

Holotype  £,  Victoria:  Ferntree  Gully,  x.1921  (Burns)  (NMV). 

Paratypes.  Queensland:  1  cf ,  Lamington  N.P. ,  xi.1961  (Common  &  Upton)  (ANIC);  1  cf ,  Mt  Glorious, 
xii  (TC);  2  cf ,  Mt  Nebo,  500  m,  iii  (TC). 

Certonotus  leeuwinensis  Turner  comb.  rev. 
(Figs  49,  52) 

Certonotus  leeuwinensis  Turner,  1919:  551.  Holotype  cf ,  WESTERN  AUSTRALIA  (BMNH)  [examined]. 
Asperellus  leeuwinensis  (Turner)  Townes  et  al. ,  1961: 115. 

Female.  Medium-sized  species,  fore  wing  length  6-7  mm.  Labium  with  glossae  unspecialized.  Lower  face 
at  narrowest  point  1-2-1-3  times  as  broad  as  high;  malar  space  1-1-1-2  times  as  long  as  basal  mandibular 
width.  Occipital  carina  ventrally  weak,  only  slightly  longer  than  abscissa  of  hypostomal  carina  between  it 
and  mandibular  base.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  weakly  convex,  in  dorsal 
view  slightly  protruding  from  mesoscutal  margin;  subalar  prominence  weakly  convex.  Scutellum  sparsely 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  129 

punctate,  with  strong  transverse  keel;  metapleuron  closely,  quite  finely  punctate;  submetapleural  carina 
anteriorly  expanded  into  a  moderately  broad,  triangular  lobe;  metanotum  with  weak  lateral  tooth. 
Propodeum  quite  short  with  anterior  transverse  carina  complete;  posterior  transverse  carina  absent; 
lateromedian  longitudinal  carina  present  before  anterior  transverse  carina;  pleural  carina  more  or  less 
complete;  area  superomedia  undefined;  area  spiracularis  incomplete  internally;  first  and  second  lateral 
areae  confluent.  Fore  wing  with  3r-m  convergent  towards  2r-m  in  some  specimens;  2m-cu  joining  M  just 
basad  of  2r-m.  Hind  wing  with  distal  abscissa  of  Cui  absent.  Hind  tibia  with  posterior  margin  with  one 
spine-like  bristle.  Caster  with  segment  1  short,  the  sternite  0-5  times  as  long  as  hind  coxa,  reaching  nearly 
to  level  of  spiracle.  Tergite  7  mediodorsally  with  indentation  to  0-5  its  length;  tergite  8  posteriorly  short  and 
rounded;  tergite  9  in  dorsal  view  short  and  broad.  Ovipositor  projecting  beyond  apex  of  gaster  by  5-5  times 
length  of  hind  tibia. 

Coloration.  Antenna  black  with  apical  segments  white  (except  for  distal  apex  of  last  segment  which  is 
black);  head  black,  orbits  entirely  whitish,  lower  face  centrally  brownish;  alitrunk  reddish  brown,  the 
sclerites  often  margined  with  black  irregularly;  pronotum  dorsally  as  a  stripe,  ventrally,  tegula,  subalar 
prominence,  scutellum  and  postscutellum  pale  whitish  yellow;  gaster  brownish  or  even  blackish,  tergites 
1-7  laterally  and  posteriorly  margined  with  yellow.  Legs  with  fore  and  mid  coxae  yellow  above,  black 
below;  hind  coxa  brown  above,  blackish  below;  anterior  two  pairs  of  legs  otherwise  brownish  except  for 
distal  apices  of  femora  which  are  yellow;  hind  femur  brownish;  hind  tibia  darker  brown,  proximally 
indistinctly  paler,  tarsus  infuscate.  Pterostigma  dark  brown,  wings  hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  3-6  mm;  malar  space  0-9  times  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  moderately  stout,  the  sternite  0-6  times  as  long  as  hind  coxa;  apex  of 
gonosquama  flattened  slightly  and  bearing  long  scattered  hairs.  Colour  similar  to  female  but  lower  face  and 
anterior  two  pairs  of  coxae  almost  entirely  yellow. 

REMARKS.  Certonotus  leeuwinensis  belongs  to  the  leeuwinensis-group.  It  is  structurally  most  similar  to  C. 
toolangi  in  having  a  noticeably  punctate  metapleuron.  The  most  obvious  difference  between  the  two 
species  is  in  coloration.  The  gaster  of  leeuwinensis  has  the  hind  margins  of  tergites  2-5  banded  with  yellow 
whereas  those  of  toolangi  are  spotted  with  yellow.  The  hind  tibia  of  toolangi  is  bicoloured,  that  of 
leeuwinensis  is  almost  unicolorous.  C.  leeuwinensis  has  a  slightly  longer  ovipositor  and  malar  space  than 
toolangi. 

MATERIAL  EXAMINED 

Holotype  cf ,  Western  Australia:  Yallingup  near  Cape  Naturaliste  ix-x.1913  (Turner)  (BMNH). 

New  South  Wales:  1  $,  Killara,  xii.1935  (Day)  (ANIC).  Tasmania:  1  $,  Coles  Bay,  ii-iii  (TC);  1  cf, 
Georgetown,  xi.1917  (Cole)  (NMV);  1  $,  Mt  Barrow,  1200  m,  xiii-i  (TC).  Western  Australia:  1  $ 
(paratype),  same  data  as  holotype  (BMNH);  1  cf ,  21  km  SW.  by  S.  Donnybrook  (23-44S  115-41E),  x.1981 
(Naumann  &  Cardale)  (ANIC);  1  cf ,  Mt  Chudalup,  S.  of  Northcliffe,  x.1970  (Colless)  (ANIC);  1  cf , 
Yallingup,  near  Cape  Naturaliste,  ix-x.1913  (Turner)  (BMNH). 

Certonotus  mogimbensis  Cheesman  comb.  rev. 
(Fig.  47) 

Certonotus  mogimbensis  Cheesman,  1936:  180.  Holotype  $,  NEW  HEBRIDES  (BMNH)  [examined]. 
Asperellus  mogimbensis  (Cheesman)  Townesefa/.,  1961: 115. 

Female.  Small  species,  fore  wing  length  5  mm.  Labium  with  glossae  slightly  elongate.  Lower  face  at 
narrowest  point  1-0  times  as  broad  as  high;  malar  space  0-8-1-0  times  as  long  as  basal  mandibular  width. 
Occipital  carina  ventrally  from  slightly  shorter  to  slightly  longer  than  abscissa  of  hypostomal  carina 
between  it  and  base  of  mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  weakly  convex, 
in  dorsal  view  parallel  to  mesoscutal  margin;  subalar  prominence  very  weakly  convex.  Scutellum  punctate 
with  strong  transverse  keel;  metapleuron  smooth  with  some  fine  punctures  posteriorly;  submetapleural 
carina  anteriorly  expanded  into  a  rounded  lobe;  metanotum  without  a  lateral  tooth.  Propodeum  short  with 
anterior  transverse  carina  complete;  posterior  transverse  carina  absent;  lateromedian  longitudinal  carina 
vestigial;  pleural  carina  complete;  area  superomedia  undefined;  area  spiracularis  not  defined  internally; 
first  and  second  lateral  areae  confluent.  Fore  wing  with  3r-m  present  for  half  the  distance  or  less  from  Rs 
towards  M;  2m-cu  joining  M  well  distad  of  2r-m.  Hind  wing  with  distal  abscissa  of  Cui  absent.  Hind  tibia 
with  posterior  margin  with  one  or  two  spine-like  bristles.  Gaster  with  segment  1  short  and  broad,  the 
sternite  0-5  times  as  long  as  hind  coxa,  reaching  to  level  of  spiracle.  Tergite  7  mediodorsally  with 
indentation  to  0-5  of  length;  tergite  8  posteriorly  long  and  rounded;  tergite  9  in  dorsal  view  elongate. 
Ovipositor  projecting  beyond  apex  of  gaster  by  4-0-4-5  times  length  of  hind  tibia. 


130  I.   D.  GAULD  &  G.  A.  HOLLOWAY 

Coloration.  Antenna  black  with  a  subapical  broad  white  band;  head  whitish;  clypeus  and  mouth  parts 
brownish,  frons  centrally  and  interocellar  area  dark  brown.  Alitrunk,  legs  and  gaster  uniformly  orange. 
Pterostigma  dark  brown,  wings  hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  3-4  mm;  malar  space  0-9-1-1  times  basal  mandibular 
width;  3r-m  absent;  gaster  with  segment  1  moderately  slender;  the  sternite  0-6  times  as  long  as  hind  coxa; 
apex  of  gonosquama  with  small  lobe  bearing  fine  scattered  hairs.  Similarly  coloured  to  female  but  with 
head  uniformly  orange,  flagellum  entirely  black,  hind  tibia  and  tarsus  weakly  to  strongly  infuscate,  some 
tergites  of  gaster  infuscate  and  wings  slightly  infumate. 

REMARKS.  C.  mogimbensis  most  closely  resembles  C.  pineus  in  colour  and  structure  and  the  two  may  be 
closely  related.  However,  mogimbensis  is  distinctive,  not  only  in  having  a  flatter  pronotum  but  in  having 
3r-m  incomplete  in  the  male  and  lacking  dark  maculae  on  the  mesoscutum.  This  species  is  known  to  occur 
in  tropical  Queensland  and  on  Vanuatu  (New  Hebrides). 

MATERIAL  EXAMINED 

Holotype  $,  New  Hebrides:  Malekula,  Ounua,  ii.1929  (Cheesman)  (BMNH). 

Queensland:  1  $,  1  cf,  Claudie  R,  near  Mt  Lamond,  xii.1971  (McAlpine  &  Holloway)  (AM);  1  cf, 
Shipton's  Flat  (15-47S  124-14E),  x.1980  (Cardale)  (ANIC). 

Certonotus  monticola  Morley 

(Fig.  38) 
Certonotus  monticola  Morley,  1913:  29.  Holotype  $,  QUEENSLAND  (BMNH)  [examined]. 

Female.  Medium-sized  species,  fore  wing  length  9-12  mm.  Labium  with  glossae  elongate,  projecting 
beyond  clypeus  by  a  distance  equal  to  or  greater  than  facial  height.  Lower  face  at  narrowest  point  0-9-1-0 
times  as  broad  as  high;  malar  space  0-5-0-6  times  as  long  as  basal  mandibular  width.  Occipital  carina 
ventrally  obsolescent,  represented  by  a  stub  that  is  shorter  than  abscissa  of  hypostomal  carina  between  it 
and  base  of  mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  weakly  convex,  in  dorsal 
view  barely  projecting  beyond  scutal  margin;  subalar  prominence  very  weakly  convex.  Scutellum  closely 
and  coarsely  punctate,  with  transverse  crest;  metapleuron  anteriorly  punctate,  posteriorly  and  dorsally; 
submetapleural  carina  anteriorly  abruptly  expanded  into  a  broad  rounded  lobe;  metanotum  without  a 
tooth  before  lateral  carina.  Propodeum  quite  short  with  anterior  transverse  carina  centrally  incomplete; 
posterior  transverse  carina  absent;  lateromedian  longitudinal  carina  present  only  before  anterior  carina; 
pleural  carina  complete;  area  superomedia  not  delineated;  area  spiracularis  complete,  short;  first  and 
second  lateral  areae  confluent.  Fore  wing  with  3r-m  converging  on  2r-m  but  joining  Rs  separately;  2m-cu 
joining  M  midway  between  3r-m  and  2r-m.  Hind  wing  with  distal  abscissa  of  Cu^  distinct  to  wing  margin. 
Hind  tibia  with  posterior  margin  without  spine-like  bristles.  Gaster  with  segment  1  short  and  broad,  the 
sternite  0-7  times  as  long  as  hind  coxa,  reaching  just  beyond  level  of  spiracle.  Tergite  7  mediodorsally  with 
small  indentation;  tergite  8  posteriorly  elongate  and  narrow;  tergite  9  in  dorsal  view  long  and  truncate. 
Ovipositor  projecting  beyond  apex  of  gaster  by  4-5  times  length  of  hind  tibia. 

Coloration.  A  reddish  brown  species  with  lower  face,  frontal  and  genal  orbits,  central  mesoscutal  spot, 
scutellum,  most  of  pronotum,  much  of  mesopleuron  and  virtually  all  of  metapleuron/propodeum  behind 
spiracle,  yellow.  Gaster  dark  brown,  tergites  2+  with  anterolateral  triangular  marks.  Fore  leg  brownish 
yellow,  coxa  infuscate;  mid  leg  blackish  brown,  femur  distally  yellow  marked,  coxa  darker;  hind  leg  with 
coxa  black  with  dorsal  yellow  marks,  femur  and  tarsus  strongly  infuscate,  tibia  less  strongly  infuscate. 
Pterostigma  black,  wings  weakly  infumate. 

Male.  Slender  gaster,  otherwise  similar  to  female  but  with  fore  wing  length  7-11  mm;  malar  space 
0-5-0-6  times  basal  mandibular  width;  3r-m  present;  gaster  exceptionally  elongate  with  segment  1  quite 
stout,  the  sternite  0-6  times  as  long  as  hind  coxa;  gonosquama  long,  apex  a  little  flattened,  with  scattered 
hairs  of  various  length.  Male  similar  in  general  colour  pattern  to  female,  although  with  fewer  yellow 
maculae. 

REMARKS.  C.  monticola  belongs  to  the  /7av/ce/w-group.  It  is  the  only  taxon  in  this  complex  to  occur  in 
Australia  where  it  is  easily  recognizable  by  the  elongate  glossae  and  short  occipital  carina.  C.  monticola  is 
known  to  occur  in  both  north  Queensland  and  Papua  New  Guinea. 

MATERIAL  EXAMINED 

Holotype  9>  Queensland:  Tambourine  Mt  (BMNH). 

Queensland:  1  $,  Middle  Claudie  R.,  Iron  Range,  x.1974  (Daniels)  (AM).  Papua  New  Guinea:  1  $, 
Amuk,  165  m,  i.1960  (Mao)  (BPBM);  1  $,  2  cf ,  Bulolo,  ix.1981,  exMyristica  sp.  (Roberts)  (BMNH);  7  $, 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  131 

3  Cf ,  Kiunga,  Fly  River,  viii-x.  1957  (Brandt)  (BPBM) ;  2  £ ,  Normanby  Is. ,  Wakaiuma,  Sewa  Bay,  xii.  1956 
(Brandt)  (BPBM);  1  $,  10  mi.  W.  Vudal,  xi.1970  (TC);  1  $,  Waris,  S.  of  Hollandia,  450-500  m,  viii.1959 
(Maa)  (BPBM). 

Certonotus  nitidulus  Morley 
(Fig.  33) 

Certonotus  nitidulus  Morley,  1913:  29.  Holotype  cf ,  VICTORIA  (BMNH)  [examined]. 
Certonotus  tasmaniensis  Turner,  1919:  550;  Hocking,  1967:  57;  Short,  1978:  41.  Holotype  cf ,  TASMANIA 
(BMNH)  [examined].  Syn.  n. 

Female.  Medium  to  large  species,  fore  wing  length  7-14  mm.  Labium  with  glossae  unspecialized.  Lower 
face  at  narrowest  point  1-0-1-1  times  as  broad  as  high;  malar  space  0-6-0-7  times  as  long  as  basal 
mandibular  width.  Occipital  carina  ventrally  strong,  slightly  longer  than  abscissa  of  hypostomal  carina 
between  it  and  base  of  mandible .  Upper  part  of  pronotum ,  slightly  before  posterior  corner ,  weakly  convex , 
in  dorsal  view  barely  projecting  beyond  scutal  margin;  subalar  prominence  weakly  convex.  Scutellum 
punctate,  transverse  keel  strong;  metapleuron  smooth  with  isolated  punctures;  submetapleural  carina 
anteriorly  expanded  into  a  broad  rounded  lobe;  metanotum  with  a  weak  tooth  opposite  anterior  end  of 
lateral  carina.  Propodeum  quite  short  with  anterior  transverse  carina  centrally  incomplete;  posterior 
transverse  carina  absent;  lateromedian  longitudinal  carina  present  only  before  anterior  carina;  pleural 
carina  present,  complete  but  rather  weak;  area  superomedia  undefined;  area  spiracularis  complete, 
defined  posteriorly  by  an  arched  carina  that  is  very  close  to  spiracle;  first  and  second  lateral  areae 
separated,  distally  delineated.  Fore  wing  with  3r-m  converging  towards  2r-m,  joining  Rs  separately;  2m-cu 
joining  M  to  0-3  basad  of  3r-m.  Hind  wing  with  distal  abscissa  of  Cu\  distinct  to  wing  margin.  Hind  tibia  with 
posterior  margin  without  spine-like  bristles.  Gaster  with  segment  1  short,  the  sternite  0-6  times  as  long  as 
hind  coxa,  reaching  behind  level  of  spiracle.  Tergite  7  mediodorsally  with  indentation  almost  to  0-5  length 
of  tergite;  tergite  8  posteriorly  short  and  rounded;  tergite  9  in  dorsal  view  transverse.  Ovipositor  projecting 
beyond  apex  of  gaster  by  5-0  times  length  of  hind  tibia. 

Coloration.  Colour  somewhat  variable  but  generally  with  antenna  black,  distally,  except  for  apex  of  last 
segment,  white.  Head  red-brown,  lower  face  laterally  to  entirely,  genal  and  frontal  orbits  pale  yellow. 
Alitrunk  reddish  brown,  yellow  marks  on  pronotum  medially,  anterior  'corners'  and  central  spot  on 
mesoscutum,  scutellum  laterally,  tegula,  subalar  prominence,  anterior  and  posterior  mesopleural  spots, 
postscutellum  and  hind  part  of  propodeum.  Gaster  reddish  with  tergites  posteriorly  and  laterally  yellowish 
margined,  and  also  with  spots  in  anterolateral  corners.  Fore  leg  yellow,  femur  red  centrally,  peripherally 
black  with  distal  apex  yellow.  Mid  leg  similar  with  coxa  black-marked  and  tibia  and  tarsus  infuscate.  Hind 
leg  as  mid  leg  but  with  coxa  reddish  brown,  ventrally  black.  Pterostigma  black,  wings  hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  7-10  mm;  malar  space  0-4-0-6  times  basal  mandibular 
width;  3r-m  present;  gaster  with  segment  1  moderately  slender,  0-6  times  as  long  as  hind  coxa;  apex  of 
gonosquama  flattened,  bearing  two  widely  separated  dense  tufts  of  long  hairs.  Similarly  coloured  to 
female. 

REMARKS.  C.  nitidulus  is  apparently  one  of  the  commonest  species  in  the  genus  as  it  has  adapted  to 
parasitizing  an  introduced  siricid  in  Pinus  radiata  plantations  in  Victoria  and  Tasmania  (Hocking,  1967).  In 
coloration  C.  nitidulus  most  strongly  resembles  C.  geniculatus  (from  which  it  can  be  separated  by  reference 
to  couplet  8  in  the  key)  but  structurally  it  is  most  similar  to  C.  annulatus  and  C.  andrewi.  C.  nitidulus  differs 
from  both  in  colour  pattern  and  in  the  form  of  the  propodeal  carinae.  This  is  a  quite  widespread  species  in 
the  south-east,  extending  from  southern  Queensland  to  Tasmania. 

HOST  RECORDS.  Siricidae:  SirexnoctilioF.  (Hocking,  1967). 

MATERIAL  EXAMINED 

Holotype  cf  (Gertonotus  nitidulus  Morley),  Victoria  (BMNH).  Holotype  cf  (Certonotus  tasmaniensis 
Turner),  Tasmania:  Mt  Wellington,  i-ii.1913  (Turner)  (BMNH). 

Queensland:  1  $ ,  Mt  Glorious,  xi  (TC);  1  cf ,  Mt  Norman  area,  Wallangarra,  x.1972  (Monteith)  (ANIC). 
New  South  Wales:  1  $ ,  Barrington  Nat.  Park,  i  (TC);  1  $ ,  Boyd  River  crossing,  Kanangra-Boyd  Nat.  Park, 
xii.  1977  (Daniels)  (AM);  1  cf ,  Leather  Barrel  Ck,  Kosciusko,  xi.1961  (Colless)  (ANIC);  1  $,  Lord  Howe 
Is.,  ii-iii.1957  (Leipa)  (BPBM);  1  cf ,  Monga,  x.1957  (Riek)  (ANIC);  1  $,  Moonee,  xi.1947  (NMV);  1  cf , 
Moss  Vale,  xi.1919  (Duquef)  (AM);  1  $,  2  cf,  Mt  Tomah,  xi.1983  (Rodd)  (AM);  1  cf,  Pebbly  Beach, 
xi.1960  (Common  &  Upton)  (ANIC);  1  cf , Tubrabucca,  xi.1953  (Burns)  (NMV).  Victoria:  1  $,  Dynamite 
Ck,  Bonang  Hwy,  x.1961  (Colless)  (ANIC);  1  $,  Harrietville,  i.1924  (Oke)  (NMV);  3  cf,  Healesville, 


132  I.  D.  GAULD  &  G.  A.  HOLLOWAY 

xi.1943  (Oke)  (NMV);  2  $ ,  1  cf ,  South  Melbourne,  breeding  cages,  x.1969,  ex  Sirex  sp.  (Waugh)  (NMV); 
3  $ ,  3  Cf ,  Mirboo  North,  x.1967,  ex  Sirex  noctilio  (Elliott)  (ANIC);  3  $ ,  Mt  Dandenong,  300  m,  ii  (TC);  3 
Cf ,  Mt  Buffalo,  1600  m,  ii  (TC);  1  $,  1  cf ,  Noorimbee,  xi.1965  (Neboiss)  (NMV);  5  cf ,  Toolangi,  xii.1982 
(Farrugia)  (BMNH) ;  5  $ ,  Toolangi ,  i-ii .  1983  (Farrugia  &  Gauld)  (BMNH) ;  2  $ ,  Warburton ,  ii-iii  (TC) ;  8 
$,  2  cf ,  Vic.  Dept  Agric.,  ii.1968,  ex  Finns  logs  (Irvine)  (ANIC).  Tasmania:  1  $,  Harrison  Ck,  between 
Cracroft  and  Blakes  Opening,  ii.1966  (Neboiss)  (MNV);  1  $,  E.  Blakes  Opening,  ii.1966  (Neboiss) 
(NMV);  1  $,  1  Cf ,  Bruny  Is.,  vii.1964,  from  Pinus  radiata  (Wilson  &  Hocking)  (AM);  1  $,  Catamaran,  ii 
(TC);  1  $,  Collinsville,  300  m,  ii.1983  (Gauld)  (BMNH);  2  $,  Frenchman's  Gap  Trig  at  Franklin  River, 
ii-iii  (TC);  1  $,  Geeveston,  ii  (TC);  1  $,  1  cf,  Hartz  Mtns,  ii-iii  (TC);  1  $,  Hellyer  Gorge,  ii.1983 
(Naumann  &  Cardale)  (ANIC);  1  $ ,  Hellyer  Gorge,  xii.1981  (Naumann  &  Cardale)  (ANIC);  2  $ ,  Hellyer 
Gorge,  ii.1967  (Riek)  (ANIC);  1  (7,  Hellyer  Gorge,  300  m,  i-ii  (TC);  1  $ ,  Huon-Picton  Junction,  ii.1967 
(Riek)  (ANIC);  1  cf ,  King  William  Ck,  xii.1981  (Gauld)  (BMNH);  2  $ ,  1  cf ,  King  William  Range  i  (TC); 
28  $ ,  18  cf ,  Launawanna,  Bruny  Is. ,  ix-x.1964  (Hocking)  (ANIC,  BMNH,  TC);  1  $ ,  1  cf ,  Mt  Barrow,  300 
m,  xii-i(TC). 

Certonotus  paluma  sp.  n. 

(Figs  54,  62) 

Female.  Medium-sized  species,  fore  wing  length  8-10  mm.  Labium  with  glossae  unspecialized.  Lower  face 
at  narrowest  point  1-1-1-2  times  as  broad  as  high;  malar  space  0-8  times  as  long  as  basal  mandibular  width. 
Occipital  carina  ventrally  as  long  as  abscissa  of  hypostomal  carina  between  it  and  base  of  mandible.  Upper 
part  of  pronotum,  slightly  before  posterior  corner,  very  weakly  convex,  in  dorsal  view  almost  parallel  with 
mesoscutal  margin;  subalar  prominence  weakly  convex.  Scutellum  sparsely  punctate,  with  a  posterior 
transverse  crest;  metapleuron  polished,  smooth,  virtually  impunctate;  submetapleural  carina  anteriorly 
expanded  into  a  broad  rounded  lobe;  metanotum  barely  produced  opposite  anterior  margin  of  lateral 
carina.  Propodeum  quite  short  with  anterior  transverse  carina  complete;  posterior  transverse  carina 
present  at  least  centrally;  lateromedian  longitudinal  carina  present  only  anterior  to  anterior  transverse 
carina;  pleural  carina  complete  anteriorly,  posteriorly  obsolescent;  area  superomedia  not  bounded 
laterally;  area  spiracularis  completely  delineated;  first  and  second  lateral  areae  separated  by  carina.  Fore 
wing  with  3r-m  converging  towards  2r-ra,  joining  the  latter  at  Rs;  2m-cu  joining  M  at  3>r-m.  Hind  wing  with 
distal  abscissa  of  Cu\  present  over  distal  0-5  to  wing  margin,  not  joined  to  first  abscissa  of  Cu\.  Hind  tibia 
with  posterior  margin  without  spine-like  bristles.  Gaster  with  segment  1  long  and  narrow,  the  sternite  0-7 
times  as  long  as  hind  coxa,  reaching  to  level  of  spiracle.  Tergite  7  mediodorsally  with  narrow  indentation, 
over  0-5  of  its  length;  tergite  8  posteriorly  long  and  narrow;  tergite  9  in  dorsal  view  short  and  pointed. 
Ovipositor  projecting  beyond  apex  of  gaster  by  4  times  length  of  hind  tibia. 

Coloration.  Antenna  black,  distally  white  with  extreme  apex  of  last  segment  black;  head  black  with  face 
and  orbits  white;  ali trunk  black  or  reddish  black,  with  central  mesoscutal  stripe,  scutellum,  pronotum, 
tegula,  subalar  prominence,  mesopleuron  centrally,  metapleuron  and  propodeum  laterally  white;  gaster 
black  with  very  large  lateral  triangular  areas  white.  Anterior  two  pairs  of  legs  white  with  femora,  tibiae  and 
tarsi  variously  infuscate ;  hind  leg  black ,  coxa  dorsally  with  white  spot .  Pterostigma  blackish ,  wings  hyaline . 

Male.  Unknown. 

REMARKS.  C.  paluma  is  one  of  the  three  quite  closely  related  Certonotus  species  that  have  an  incomplete 
distal  abscissa  of  Cu\  in  the  hind  wing.  It  is  distinguishable  from  Certonotus  sp.  A  by  its  unspecialized 
subalar  prominence.  It  is  structurally  similar  to  C.  ixion,  from  which  it  may  be  separated  by  reference  to  the 
key. 

MATERIAL  EXAMINED 

Holotype  9,  Queensland:  Paluma,  900  m  (19-OOS  146-12E),  x.1980,  ex  Malaise  trap  (Frith)  (ANIC). 
Paratype.  Queensland:  1  $,  Windsor  Tableland,  iii.1981  (Storey)  (BMNH). 

Certonotus  pineussp.  n. 

(Figs  45,  46) 

Female.  Medium-sized  species,  fore  wing  length  7-9  mm.  Labium  with  glossae  unspecialized.  Lower  face 
at  narrowest  point  1-0-1-1  times  as  broad  as  high;  malar  space  0-9-1-0  times  as  long  as  basal  mandibular 
width.  Occipital  carina  ventrally  almost  3  times  as  long  as  abscissa  of  hypostomal  carina  between  it  and  base 
of  mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  very  convex,  in  dorsal  view 
subpyramidal;  subalar  prominence  moderately  convex.  Scutellum  moderately,  closely  punctate,  trans- 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  133 

verse  keel  weak;  metapleuron  anteriorly  smooth,  posteriorly  punctate;  submetapleural  carina  anteriorly 
expanded  into  a  broad  almost  quadrate  lobe;  metanotum  with  a  very  weak  tooth  before  vestige  of  lateral 
carina.  Propodeum  quite  short  with  anterior  transverse  carina  present,  very  close  to  anterior  part  of 
propodeum;  posterior  transverse  carina  absent;  lateromedian  longitudinal  carina  vestigial  before  anterior 
carina,  otherwise  absent;  pleural  carina  more  or  less  complete;  area  superomedia  undefined;  area 
spiracularis  more  or  less  complete  except  internally;  first  and  second  lateral  areae  not  separated.  Fore  wing 
with  3r-m  only  slightly  convergent  towards  2r-m,  joining  Rs  away  from  2r-m,  making  areolet  quadrate; 
2m-cu  joining  M  at  3r-m.  Hind  wing  with  distal  abscissa  of  Cu\  absent.  Hind  tibia  with  posterior  margin 
with  one  spine-like  bristle.  Gaster  with  segment  1  short  and  broad,  the  sternite  0-3  times  as  long  as  hind 
coxa,  just  not  reaching  level  of  spiracle.  Tergite  7  mediodorsally  with  indentation  to  0-5  its  length;  tergite  8 
posteriorly  long  and  rounded;  tergite  9  in  dorsal  view  short  and  broad.  Ovipositor  projecting  beyond  apex 
of  gaster  by  4-5  times  length  of  hind  tibia. 

Coloration.  Antenna  black,  with  subapical  white  band;  head  yellowish,  only  interocellar  area  and  edges 
of  mandibles  blackish.  Alitrunk  and  gaster  yellowish;  mesoscutum  with  lateral  marks,  anterocentrally 
infuscate;  tip  of  tergite  8  blackish.  Legs  yellow,  hind  tarsus  and  sometimes  distal  apex  of  tibia  blackish. 
Pterostigma  black,  wings  hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  4  mm;  malar  space  0-7  times  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  quite  stout,  the  sternite  0-3  times  as  long  as  hind  coxa;  apex  of 
gonosquama  rounded,  bearing  scattered  hairs.  Similar  in  colour  to  female  but  with  yellowish  areas  a  little 
paler,  more  strongly  contrasted  with  infuscate  areas;  flagellar  white  mark  very  indistinct. 

REMARKS.  A  stocky  species  easily  recognized  by  the  almost  pyramidal  pronotal  convexity.  The  virtually 
quadrate  areolet  and  very  short  first  sternite  are  also  quite  characteristic  of  this  species.  C.  pineus  is  only 
known  to  occur  in  Queensland. 

MATERIAL  EXAMINED 

Holotype  $,  Queensland:  Moses  Ck,  45  km  N.  by  E.  Mt  Finnigan  (15-47S  145-17E),  x.1980  (Cardale) 
(ANIC). 

Paratypes.  Queensland:  1  $,  Lake  Barrine,  ii.1935  (Burns)  (NMV);  1  cf ,  Mt  Webb  Nat.  Park  (15-04S 
145-07E),  iv.1981  (Naumann)  (ANIC);  1  cf ,  Palm  Is.  nearTownsville,  x  (TC). 

Certonotus  rufescens  Morley 
(Figs  22,  25) 

Certonotus  rufescens  Morley,  1913:  30.  LECTOTYPE  $,  QUEENSLAND  (BMNH),  here  designated 
[examined] . 

Female.  Moderately  large  to  large  species,  fore  wing  length  8-16  mm.  Labium  with  glossae  slightly 
lengthened.  Lower  face  at  narrowest  point  0-9-1  -0  times  as  broad  as  high;  malar  space  0-9-1-0  times  as  long 
as  basal  mandibular  width.  Occipital  carina  ventrally  rather  straight,  more  than  twice  as  long  as  abscissa  of 
hypostomal  carina  between  it  and  mandible  base.  Upper  part  of  pronotum,  slightly  before  posterior 
corner,  weakly  convex,  in  dorsal  view  rounded,  just  protruding  beyond  scutal  margin;  subalar  prominence 
very  strongly  raised,  in  dorsal  view  with  a  blunt,  back-curved,  thorn-like  protuberance.  Scutellum 
punctate,  with  a  moderately  strong  transverse  crest;  metapleuron  smooth,  virtually  impunctate;  submeta- 
pleural carina  anteriorly  expanded  into  a  broad  almost  quadrate  lobe  which  usually  has  ridge  near  anterior 
corner  delimiting  a  narrow  triangular  area;  metanotum  with  small  tooth  opposite  anterior  end  of  lateral 
carina.  Propodeum  moderately  long  with  anterior  transverse  carina  complete;  posterior  transverse  carina 
present  as  lateral  vestige;  lateromedian  longitudinal  carina  present  before  anterior  carina;  pleural  carina 
very  strong  anteriorly,  absent  behind  anterior  carina;  area  superomedia  not  delineated  laterally;  area 
spiracularis  complete;  first  and  second  lateral  areae  confluent,  not  delineated  laterally.  Fore  wing  with 
3r-m  converging  towards  2r-m  and  joining  latter  at  Rs;  2m-cu  joining  M  at  3r-m  to  0-3  towards  2r-m.  Hind 
wing  with  distal  abscissa  of  Cui  distinct  to  wing  margin.  Hind  tibia  with  posterior  margin  with  one  to  three 
spine-like  bristles.  Gaster  with  segment  1  long  and  narrow,  the  sternite  0-9-1-0  times  as  long  as  hind  coxa, 
reaching  behind  level  of  spiracle.  Tergite  7  mediodorsally  at  most  only  slightly  indented  on  posterior 
margin;  tergite  8  posteriorly  with  process  short  and  rounded;  tergite  9  in  dorsal  view  elongated  and  bluntly 
pointed  posteriorly.  Ovipositor  projecting  beyond  apex  of  gaster  by  4-5  times  length  of  hind  tibia. 

Coloration.  Brownish  orange;  flagellum  black  with  a  white  subapical  band  which  is  usually  well 
developed  but  is  entirely  absent  in  specimens  from  the  northern  part  of  the  range.  Pterostigma  dark  brown, 
wings  hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  5-7  mm;  malar  space  0-8-0-9  times  basal  mandibular 


134  I.  D.  GAULD  &  G.  A.  HOLLO  WAY 

width;  3r-m  present;  gaster  with  segment  1  long,  slender,  the  sternite  0-8-1-0  times  as  long  as  hind  coxa; 
apex  of  gonosquama  simply  truncate,  not  broadened,  but  with  long  sparse  hairs.  Similarly  coloured  to 
female,  flagellum  without  a  white  band. 

REMARKS.  C.  rufescens  is  easily  distinguished  from  other  Australian  species  by  the  thorn-like  subalar 
prominence.  The  face  of  this  species  is  flatter  than  most  other  Certonotus  and  laterally  it  is  abruptly,  almost 
angularly  rounded  before  the  malar  space.  The  characteristic  submetapleural  carina  and  possession  of  a 
strong,  usually  Y-shaped  vestigial  carina  above  the  insertion  of  the  hind  coxa  are  useful  confirmatory 
characters.  A  widespread  species  extending  from  northern  Queensland  south  to  near  Melbourne,  Victoria. 

MATERIAL  EXAMINED 

Lectotype  9,  Queensland:  Mackay,  ix.1901  (BMNH). 

Queensland:  2  $  (paralectotypes),  same  data  as  lectotype  (BMNH);  2  cf ,  Mackay,  1909  (BMNH);  1  $ , 
Montville,  ix.1935  (Burns)  (NMV);  1  cf ,  Moses  Ck,  4  km  N.  by  E.  Mt  Finnigan  (15-47S  145-17E),  x.1980, 
at  light  (Cardale)  (ANIC);  1  $,  Mt  Glorious,  xii.1979  (Galloway}  (BMNH);  1  $,  Mt  Glorious  near 
Brisbane,  xii.1976  (Boucek)  (BMNH);  3  cf,  Mt  Glorious,  i-iii  (TC);  2  $,  Mt  Tambourine,  ix-x.1978 
(Galloway)  (BMNH);  2  $ ,  4  cf ,  Mt  Tambourine,  x-xi.  1977  (Galloway)  (BMNH).  New  South  Wales:  1  $ , 
19  km  S.  Coff  s  Harbour,  i.1958  (Riek)  (ANIC);  1  cf ,  Otford,  xii.1962  (Colless)  (BMNH);  1  $ ,  Sassafrass 
Gully,  Springwood,  ix.1972  (McAlpine)  (AM).  Victoria:  2  $,  Toolangi,  i-ii.1983  (Farrugia  &  Gauld) 
(BMNH);  1  Cf ,  Yellingbo,  xi.1976  (Neboiss)  (NMV). 

Certonotus  sisyphussp.  n. 

(Figs  36,  39, 40) 
[Certonotus  hinnuleus  Krieger;  Morley  1913:  32.  Misidentification.] 

Female.  Medium  to  large-sized  species,  fore  wing  length  8-18  mm.  Labium  with  glossae  slightly  leng- 
thened. Lower  face  at  narrowest  point  1-1-1-3  times  as  broad  as  high;  malar  space  0-9-1-0  times  as  long  as 
basal  mandibular  width.  Occipital  carina  ventrally  sinuous,  much  longer  than  abscissa  of  hypostomal 
carina  between  it  and  mandibular  base.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  weakly 
convex,  in  dorsal  view  projecting  slightly  beyond  scutal  margin,  subalar  prominence  very  strongly  convex, 
in  dorsal  view  slightly  pyramidal.  Scutellum  sparsely  punctate,  transverse  crest  distinct;  metapleuron 
smooth  with  scattered  punctures,  on  larger  specimens  with  punctures  closer;  submetapleural  carina 
anteriorly  expanded  into  a  rectangular  flange;  metanotum  with  strong  tooth  opposite  end  of  lateral  carina. 
Propodeum  moderately  long  with  anterior  transverse  carina  complete;  posterior  transverse  carina  strong, 
complete,  almost  parallel  to  anterior  carina;  lateromedian  longitudinal  carina  present  only  as  vestiges 
before  anterior  carina;  pleural  carina  weak,  posteriorly  evanescent;  area  superomedia  not  defined;  area 
spiracularis  complete;  first  and  second  lateral  areae  clearly  separated  though  latter  weakly  defined 
externally.  Fore  wing  with  3r-m  strongly  converging  towards  an  almost  vertical  2r-m,  joining  Rs  separately; 
2m-cu  joining  M  0-2  from  3r-m  towards  2r-m.  Hind  wing  with  distal  abscissa  of  Cui  distinct  to  wing  margin. 
Hind  tibia  with  posterior  margin  with  one  to  three  spine-like  bristles.  Gaster  with  segment  1  long  and 
narrow,  the  sternite  1-0-1-2  times  as  long  as  hind  coxa,  reaching  behind  level  of  spiracle.  Tergite  7 
mediodorsally  with  wide  indentation  to  0-5  length  of  tergite;  tergite  8  posteriorly  short  and  rounded;  tergite 
9  in  dorsal  view  as  long  as  wide  with  blunt  point  posteriorly.  Ovipositor  projecting  beyond  apex  of  gaster  by 
4-5  times  length  of  hind  tibia. 

Coloration.  Reddish  brown,  flagellum  darker  with  subapical  white  band;  clypeus,  facial,  frontal  and 
genal  orbits.  Subalar  prominence,  pair  of  central  stripes  on  mesoscutum,  scutellum,  mesopleural  spot, 
spots  in  posterolateral  corners  of  tergites  1+  and  also  on  lateral  margins  of  3+  yellow.  Legs  yellowish 
brown,  anterior  two  pairs  of  coxae  and  distal  apex  of  mid  femur  yellow.  Pterostigma  dark  brown,  wings 
hyaline. 

Male.  Similar  to  female  but  with  fore  wing  length  8  mm;  malar  space  0-7  times  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  slender,  the  sternite  1-1  times  as  long  as  hind  coxa;  apex  of 
gonosquama  with  a  small  angulate  lobe  that  bears  scattered  long  hairs.  Similar  in  colour  to  female  but  with 
face  entirely  yellow,  mid  tibia  yellow  and  yellow  maculae  on  gaster  virtually  absent. 

REMARKS.  The  fairly  elongate  petiole,  rather  flat  face  and  posteriorly  flanged  lateral  propodeal  carina 
suggest  this  species  is  related  to  C.  rufescens  though  the  pleural  carina  is  not  tuberculate  as  it  is  in  rufescens 
and  geniculatus .  Superficially  C.  sisyphus  is  similar  to  C.  avitus  from  which  it  can  be  distinguished,  not  only 
by  the  characters  in  the  key,  but  also  in  having  a  distinctly  longer  prementum  and  having  the  hypostomal 
and  occipital  carinae  meeting  further  from  the  base  of  the  mandible  than  the  basal  mandibular  width. 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  135 

Widely  distributed  in  the  south-east  of  Australia;  recorded  from  New  South  Wales,  Victoria  and 
Tasmania. 

MATERIAL  EXAMINED 

Holotype  $,  Victoria:  Toolangi,  xi-xii.1982  (Farrugia  &  Gauld)  (AM). 

Paratypes.  Victoria:  8  $ ,  1  C?,  same  data  as  holotype  (BMNH).  New  South  Wales:  2  $ ,  Acacia  Plateau, 
3000  ft,  x.1961  (Common  &  Upton)  (ANIC).  Tasmania:  1  $,  Duck  River,  6  km  SE.  Roger  R.,  xii.1974 
(Neboiss)  (NMV);  1  $,  MeridithR.,  20  km  from  Corinna,  i.  1954  (Campbell)  (ANIC);  1  $,  Mt  Field  Nat. 
Park,  250  m,  i-ii.1983  (Gauld)  (BMNH);  1  $,  Picton  R.  bridge,  i.1983  (Gauld)  (BMNH);  1  ?,  no  data 
(ANIC). 

Certonotus  talus  sp.  n. 

(Figs  34,  35) 

Female.  Medium-sized  species,  fore  wing  length  8-12  mm.  Labium  with  glossae  unspecialized.  Lower  face 
at  narrowest  point  1-1  times  as  broad  as  high;  malar  space  0-7-0-8  times  as  long  as  basal  mandibular  width. 
Occipital  carina  ventrally  more  than  twice  as  long  as  abscissa  of  hypostomal  carina  between  it  and  base  of 
mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  very  weakly  convex,  in  dorsal  view 
barely  projecting  beyond  scutal  margin;  subalar  prominence  convex,  with  a  long  sharp  back-curved  spine. 
Scutellum  punctate  with  strong  transverse  keel;  metapleuron  very  sparsely  punctate;  submetapleural 
carina  anteriorly  expanded  into  a  broad  triangular  lobe;  metanotum  with  a  large  tooth  before  anterior  end 
of  lateral  carina.  Propodeum  moderately  long  with  anterior  transverse  carina  complete;  posterior 
transverse  carina  almost  complete;  lateromedian  longitudinal  carina  vestigial,  discernible  only  before 
anterior  carina;  pleural  carina  strong  anteriorly,  posteriorly  poorly  developed  or  obsolescent;  area 
superomedia  not  delineated;  area  spiracularis  complete,  posteriorly  very  weakly  delineated;  first  and 
second  lateral  areae  present,  separated  by  strong  carina.  Fore  wing  with  3r-m  converging  towards  2r-m, 
joining  latter  at  Rs;  2m-cu  joining  M  at  3r-m.  Hind  wing  with  distal  abscissa  of  Cu\  distinct  to  wing  margin. 
Hind  tibia  with  posterior  margin  with  or  without  one  spine-like  bristle.  Gaster  with  segment  1  short,  the 
sternite  0-8-1-0  times  as  long  as  hind  coxa,  reaching  behind  level  of  spiracle.  Tergite  7  mediodorsally  with 
wide  indentation  almost  to  0-7  length  of  tergite;  tergite  8  posteriorly  with  long,  narrow  process;  tergite  9  in 
dorsal  view  long  and  evenly  rounded .  Ovipositor  projecting  beyond  apex  of  gaster  by  5  times  length  of  hind 
tibia. 

Coloration.  Antenna  black  with  subapical  white  band;  head  reddish  brown,  orbits  and  clypeus  entirely 
pale;  alitrunk  reddish,  mesoscutum  with  longitudinal  stripes,  tegula,  subalar  prominence,  scutellum 
partly,  postscutellum,  anterior  mesopleural  spot  and  paired  spots  on  hind  edge  of  propodeum  yellow; 
gaster  reddish,  lateral  margins  of  all  tergites  yellow  and  tergites  also  with  paired  large  yellow  spots  in 
posterolateral  corners.  Fore  and  mid  legs  yellow,  femur  for  proximal  0-7  and  tibia  externally  reddish,  distal 
tarsal  segment  blackish;  hind  leg  virtually  entirely  reddish  brown,  tarsus  infuscate.  Pterostigma  brown, 
wings  hyaline. 

Male.  Unknown. 

REMARKS.  The  possession  of  a  long,  backwardly  curved,  slender  spine  arising  from  the  subalar  prominence 
immediately  distinguishes  talus  from  all  other  Australian  Certonotus.  It  appears  to  belong  to  the 
humeralifer- group  but  differs  in  having  a  more  slender  petiole  with  a  much  longer  sternite.  One  specimen 
has  asymmetric  hind  wings;  the  left  wing  has  the  distal  abscissa  of  Cu\  incomplete,  not  joining  Cu±  and  cu-a, 
whilst  in  the  right  wing  the  vein  is  complete.  This  incomplete  condition  is  found  in  C.  ixion  andpaluma  and 
we  believe  these  are  closely  related  species.  All  have  similarly  modified  posterior  gastral  tergites,  and 
rather  small,  oblique  areolets.  The  petiole  is  more  slender  in  these  species  than  in  other  taxa  in  this  group 
and  the  sternite  is  longer. 

C.  talus  occurs  in  New  South  Wales  and  Victoria  and  has  been  collected  in  subtropical  and  temperate  wet 
forest. 

MATERIAL  EXAMINED 

Holotype  $,  New  South  Wales:  Dorrigo  Nat.  Park,  E.  end  of  Blackbutt  Track,  710  m,  ii-iii.1980,  in 
subtropical  rainforest  (Newtown  &  Thayer)  (ANIC). 

Paratypes.  Victoria:  2  <J>,  Toolangi,  xi.1982  (Farrugia)  (BMNH).  New  South  Wales:  1  <j>,  Tubrabucca, 
i.  1948  (Bums)  (NMV). 


136  I.  D.  GAULD  &  G.  A.  HOLLOW  AY 

Certonotus  toolangi  sp.  n. 

(Fig.  60) 

Female.  Medium-sized  species,  fore  wing  length  6-7  mm.  Labium  with  glossae  unspecialized.  Lower  face 
at  narrowest  point  1-1  times  as  broad  as  high;  malar  space  0-9-1-1  times  as  long  as  basal  mandibular  width. 
Occipital  carina  ventrally  as  long  as  basal  mandibular  width.  Occipital  carina  ventrally  as  long  as  or  slightly 
longer  than  abscissa  of  hypostomal  carina  between  it  and  base  of  mandible.  Upper  part  of  pronotum, 
slightly  before  posterior  corner,  almost  flat,  in  dorsal  view  parallel  to  mesoscutal  margin;  subalar 
prominence  moderately  convex.  Scutellum  sparsely  punctate,  transverse  keel  weak;  metapleuron  with 
shallow  coarse  close  punctures;  submetapleural  carina  anteriorly  expanded  into  a  broad  triangular  lobe; 
metanotum  with  a  weak  tooth  laterally.  Propodeum  quite  short  with  anterior  transverse  carina  complete; 
posterior  transverse  carina  absent;  lateromedian  longitudinal  carina  vestigial,  only  present  before  anterior 
transverse  carina;  pleural  carina  present  anteriorly;  area  superomedia  undefined;  area  spiracularis 
incompletely  defined  internally;  first  and  second  lateral  areae  confluent,  undefined  laterally.  Fore  wing 
with  3r-m  convergent  towards  2r-m,  joining  Rs  separately;  2m-cu  joining  M  very  near  3r-m.  Hind  wing  with 
distal  abscissa  of  Cui  absent.  Hind  tibia  with  posterior  margin  with  one  spine-like  bristle.  Gaster  with 
segment  1  short  and  broad,  the  sternite  0-6  times  as  long  as  hind  coxa,  just  reaching  level  of  spiracle. 
Tergite  7  mediodorsally  with  indentation  to  0-5  of  length;  tergite  8  posteriorly  short  and  rounded;  tergite  9 
in  dorsal  view  short  and  broad.  Ovipositor  projecting  beyond  apex  of  gaster  by  4-0-4-5  times  length  of  hind 
tibia. 

Coloration.  Very  similarly  coloured  to  leeuwinensis  but  differing  in  having  the  anterior  two  pairs  of 
coxae  entirely  pale  and  the  tibiae  with  at  least  proximal  0-5  whitish  and  mesoscutum  centrally  with  a  pair  of 
short  yellowish  stripes. 

Male.  Unknown. 

REMARKS.  C.  toolangi  belongs  to  the  leeuwinensis-group.  It  is  most  readily  distinguishable  by  its  colour 
pattern.  Structurally  it  is  rather  similar  to  C.  leeuwinensis  from  which  it  may  be  distinguished  by  its  shorter 
malar  space  and  ovipositor.  It  is  only  known  from  the  south-east  of  Australia,  where  it  has  been  taken  in 
wet  sclerophyll  forest. 

MATERIAL  EXAMINED 

Holotype  $ ,  New  South  Wales:  New  England  Nat.  Park,  x.1962  (Colless)  (ANIC). 

Paratypes.  Victoria:  2  $,  Toolangi,  xi.1982  (Farrugia)  (BMNH);  1  $,  Toolangi,  xii.1982  (Farmgia  & 
Gauld)  (BMNH). 

Certonotus  zebrus  sp.  n. 

(Figs  51,  53,  59) 

Female.  Medium-sized  species,  fore  wing  length  7-8  mm.  Labium  with  glossae  unspecialized.  Lower  face 
at  narrowest  point  0-8-0-9  times  as  broad  as  high;  malar  space  0-8-0-9  times  as  long  as  basal  mandibular 
width.  Occipital  carina  ventrally  weak,  nearly  twice  as  long  as  abscissa  of  hypostomal  carina  between  it  and 
mandibular  base.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  very  weakly  convex,  in  dorsal 
view  parallel  with  mesoscutal  margin;  subalar  prominence  strongly  convex,  centrally  somewhat  produced. 
Scutellum  sparsely  punctate,  transverse  crest  quite  weak;  metapleuron  anteriorly  smooth,  posteriorly  with 
scattered  punctures;  submetapleural  carina  anteriorly  expanded  into  a  broad  rounded  lobe;  metanotum 
with  a  weak  tooth  opposite  anterior  end  of  lateral  carina.  Propodeum  short  with  anterior  transverse  carina 
complete;  posterior  transverse  carina  absent;  lateromedian  longitudinal  carina  discernible  as  traces  before 
anterior  transverse  carina;  pleural  carina  incomplete  posteriorly;  area  superomedia  not  delineated;  area 
spiracularis  virtually  complete;  first  and  second  lateral  areae  confluent,  not  defined  laterally.  Fore  wing 
with  3r-/n  converging  towards  2r-ra,  well  separated  at  Rs;  2m-cu  joining  M  0-3  from  3r-m  towards  2r-m. 
Hind  wing  with  distal  abscissa  of  Cui  absent.  Hind  tibia  with  posterior  margin  with  one  spine-like  bristle. 
Gaster  with  segment  1  long,  the  sternite  0-6  times  as  long  as  hind  coxa,  just  reaching  level  of  spiracle. 
Tergite  7  mediodorsally  with  indentation  to  0-5  its  length;  tergite  8  posteriorly  short  and  rounded 
posteriorly.  Ovipositor  projecting  beyond  apex  of  gaster  by  4  times  length  of  hind  tibia. 

Coloration.  A  black  species  that  sometimes  has  pale  yellow-white  marks  on  distal  flagellar  segments 
(except  extreme  apex  of  distal  one);  lower  face  entirely,  orbits,  most  of  pronotum,  tegula,  scutellum, 
postscutellum,  subalar  prominence,  mesopleural  stripe,  most  of  mesopleuron  and  propodeum  laterally 
and  posteriorly,  base  of  petiole,  posterior  and  lateral  margins  of  gastral  tergites  whitish.  Anterior  two  pairs 
of  legs  whitish,  femora  and  tibiae  centrally,  tarsi  partly  brownish  marked.  Hind  leg  black.  Pterostigma 
blackish,  wings  hyaline. 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  137 

Male.  Similar  to  female  but  with  fore  wing  length  3  mm;  malar  space  0-7  times  basal  mandibular  width; 
3r-m  present;  gaster  with  segment  1  quite  stout,  the  sternite  0-5  times  as  long  as  hind  coxa;  apex  of 
gonosquama  flattened  slightly  with  scattered  hairs.  Similarly  coloured  to  female  but  with  flagellum  entirely 
black  as  is  most  of  mesopleuron  and  metapleuron. 

REMARKS.  The  holotype  differs  from  the  female  paratype  in  having  entirely  black  antennae.  C.  zebrus 
belongs  to  the  leeuwinensis-group.  It  is  the  only  species  in  the  complex  which  is  virtually  entirely  black  and 
white.  Structurally  it  is  rather  unremarkable  but  it  has  a  more  strongly  raised  subalar  prominence  than  any 
other  species  in  this  group. 
It  is  only  known  from  tropical  Queensland. 

MATERIAL  EXAMINED 

Holotype  $,  Queensland:  Baldy  Mtn  Road,  via  Atherton,  vi.1981,  ex  Malaise  trap  (Brown)  (QM). 

Paratypes.  Queensland:  1  cT,  3  km  N.  by  E.  Mt  Tip  Tree  (17-02S  145-37E),  x.1980,  at  light  (Cardale) 
(ANIC);  1  $,  Windsor  Tableland  via  Mt  Carbine,  xii.1980,  ex  Malaise  trap  (BMNH). 

Certonotussp.  A 
(Figs  48,  50) 

Female.  Medium-sized  species,  fore  wing  length  7  mm.  Labium  with  glossae  unspecialized.  Lower  face  at 
narrowest  point  1-3  times  as  broad  as  high;  malar  space  1-0  times  as  long  as  basal  mandibular  width. 
Occipital  carina  ventrally  long,  about  3  times  length  of  abscissa  of  hypostomal  carina  between  it  and  base  of 
mandible.  Upper  part  of  pronotum,  slightly  before  posterior  corner,  moderately  convex,  in  dorsal  view 
projecting  beyond  mesoscutal  margin;  subalar  prominence  weakly  convex,  rather  sharp  and  ridge-like. 
Scutellum  coarsely,  closely  punctate,  transverse  keel  weak;  metapleuron  closely,  coarsely  punctate; 
submetapleural  carina  anteriorly  expanded  into  a  broad  rounded  flange,  metanotum  with  a  distinct  tooth 
near  anterior  end  of  lateral  carina.  Propodeum  moderately  long  with  anterior  transverse  carina  centrally 
strong;  lateromedian  longitudinal  carina  present  anteriorly  and  weak  between  transverse  carinae;  pleural 
carina  strong  anteriorly,  posteriorly  obsolescent;  area  superomedia  clearly  discernible,  hexagonal;  area 
spiracularis  complete;  first  and  second  lateral  areae  indistinctly  delineated,  confluent.  Fore  wing  with  3r-m 
converging  towards  2r-m,  joining  Rs  separately;  2m-cu  joining  M  opposite  3r-m.  Hind  wing  with  distal 
abscissa  of  Cu\  absent.  Hind  tibia  with  posterior  margin  without  spine-like  bristles.  Gaster  with  segment  1 
short,  the  sternite  0-2  times  as  long  as  hind  coxa,  not  reaching  level  of  spiracle.  Tergite  7  mediodorsally  with 
indentation  on  posterior  margin;  tergite  8  posteriorly  very  elongate  and  narrow;  tergite  9  in  dorsal  view 
quadrate,  rounded  posteriorly.  Ovipositor  projecting  beyond  apex  of  gaster  by  4  times  length  of  hind  tibia. 

Coloration.  Head  blackish  with  lower  face,  genal  and  frontal  orbits  yellow;  alitrunk  reddish  brown,  the 
tergites  peripherally  black  and  only  tegula  and  subalar  prominence  yellow.  Gaster  with  tergites  3+  with 
posterior  and  lateral  margins  very  narrowly  yellow.  Legs  reddish  brown,  anterior  two  pairs  of  coxae 
yellowish  marked,  hind  coxa,  trochanter,  femur  proximally  and  distally,  and  tarsus  infuscate.  Pterostigma 
blackish,  wings  hyaline. 

Male.  Unknown. 

REMARKS.  Certonotus  sp.  A  is  easily  recognized  by  its  colour  pattern  and  possession  of  very  slender  tergal 
processes.  The  area  superomedia  is  almost  regularly  hexagonal  and  strongly  delineated  except  for  the  weak 
posterolateral  sides.  Despite  lacking  any  trace  of  a  distal  abscissa  of  C«i  in  the  hind  wing  this  species 
appears  to  be  quite  closely  related  to  C.  ixion,  suggesting  it  belongs  in  the  humeralifer-group  rather  than  in 
the  leeuwinensis-group. 
It  is  known  only  from  Western  Australia. 

MATERIAL  EXAMINED 
Western  Australia:  1  $ ,  Swan  River  (BMNH). 

Tribe  POECILOCRYPTINI 

This  relatively  small  tribe  is  characterized  by  the  lack  of  an  occipital  carina  dorsally  and  the 
presence  of  a  single  bulla  in  vein  2m-cu  in  the  fore  wing.  It  is  restricted  to  Australia  where  it  is 
represented  by  three  genera,  Alaothyris,  Urancyla  and  Poecilocryptus . 

Very  little  is  known  about  the  biology  of  poecilocryptines,  but  the  available  records  suggest 
the  species  all  oviposit  into  nutritious  plant  tissue.  It  is  possible  that  the  larva  is  partially 
phytophagous,  as  it  has  rather  massive  mandibles  (Short,  1978). 


138  I.  D.  GAULD  &  G.  A.  HOLLOWAY 

Key  to  genera  of  Poecilocryptini 

1  Fore  wing  with  areolet  and  2r-m  obliterated  by  fusion  of  Rs  and  M  (Fig.  63);  gaster  strongly 

laterally  compressed;  ovipositor  very  long,  at  least  6-0  times  length  of  hind  tibia 

ALAOTHYRIS  Gauld(p.  138) 

-  Fore  wing  with  areolet  distinct,  bounded  internally  by  2r-m  (Fig.  5);  gaster  cylindrical  or 

depressed ;  ovipositor  less  than  5  -0  times  length  of  hind  tibia 2 

2  Hind  wing  with  distal  abscissa  of  Cu\  present;  propodeal  carinae  vestigial  dorsally;  ovipositor 

strongly decurved (Fig.  64) URANCYLA  Gauld(p.  141) 

-  Hind  wing  with  distal  abscissa  of  Cu\  absent;  propodeum  with  at  least  some  carinae  dorsally; 

ovipositor  more  or  less  straight POECILOCRYPTUS  Cameron  (p.  139) 

ALAOTHYRIS  Gauld 

Genus  A;  Gauld,  1983:  169. 

Alaothyris  Gauld,  1984:  93.  Type-species:  Alaothyris  elongissimus  Gauld,  by  original  designation. 

Medium-sized  species,  fore  wing  length  6  mm;  clypeus  small,  flat,  truncate;  labrum  moderately  large, 
exposed;  mandible  quite  short,  tapered,  twisted  about  20  times,  almost  evenly  bidentate;  malar  space 
slightly  less  than  basal  mandibular  width.  Occipital  carina  absent  on  dorsal  part  of  head,  ventrally  joining 
hypostomal  carina  above  base  of  mandible.  Antenna  long,  not  tapered.  Mesoscutum  polished,  almost 
smooth;  notaulus  deep  on  anterior  0-2  of  scutum,  notaular  crest  occluding  extreme  anterior  end;  scutellum 
weakly  convex,  not  laterally  carinate;  propodeum  long,  evenly  rounded  without  carinae  dorsally; 
propodeal  spiracles  circular;  gaster  inserted  at  end  of  short  propodeal  neck,  above  and  far  behind  coxal 
insertion.  Fore  tibia  with  a  small  tooth  on  outer  side;  mid  and  hind  coxae  very  elongate;  tarsal  claws  simple. 
Fore  wing  with  cu-a  proximal  to  base  of  Rs&M ;  3r-m  absent;  Rs  and  M  fused  to  obliterate  areolet  and  2r-m; 
2m-cu  with  a  single  bulla.  Hind  wing  with  distal  abscissa  of  Ct/i  absent;  basal  cell  slender;  Sc  bearing  one 
hamulus.  Gaster  very  long  and  slender,  laterally  compressed;  tergite  1  slender,  with  spiracles  a  little  behind 
centre;  sternite  1  reaching  far  behind  level  of  spiracles;  tergites  2-3  with  laterotergites  folded  under. 
Ovipositor  very  long  and  slender,  projecting  beyond  apex  of  gaster  by  more  than  6-0  times  length  of  hind 
tibia;  apex  cylindrical. 

REMARKS.  A  very  distinctive  genus  easily  recognized  by  its  slender  facies  and  characteristic  venation.  The 
systematic  position  of  this  genus  is  questionable.  The  mandible  and  elongate  structure  suggest  a 
relationship  with  the  Labenini,  but  the  position  of  the  petiolar  spiracle,  venation  and  shape  of  propodeum 
suggest  that  it  is  perhaps  more  closely  related  to  the  Poecilocryptini,  especially  Poecilocryptus.  Unlike  the 
Labenini,  Alaothyris  does  not  have  fine,  file-like  teeth  on  the  ovipositor  apex,  nor  apparently  does  it  have  a 
lobe  at  the  base  of  the  ovipositor  sheath.  A  single  species  is  known. 

Alaothyris  elongissimus  Gauld 

(Fig.  63) 
Alaothyris  elongissimus  Gauld,  1984:  94.  Holotype  9,  QUEENSLAND  (ANIC)  [examined]. 

Female.  Lower  face  slightly  elongate,  with  a  pronounced  central  tubercle;  eye  surface  finely  pubescent; 
ocelli  arranged  in  an  equilateral  triangle;  flagellum  with  about  28  segments.  Mesoscutum  polished, 
impunctate;  mesopleuron  and  metapleuron  similarly  smooth,  epicnemial  carina  dorsally  obsolescent; 
submetapleural  carina  broad  anteriorly.  Gaster  highly  polished. 

Coloration.  Predominantly  orange-brown  species  with  flagellum,  hind  leg  and  gaster  darker  brown. 
Pterostigma  brown,  wings  hyaline. 

Male.  Similar  to  female. 

REMARKS.  The  holotype  and  paratype  emerged  from  the  seeds  ofAraucaria  cunninghamii.  What  their  host 
was  is  not  known.  Probably  it  will  be  found  to  be  some  seed-feeding  beetle,  but  the  possibility  (given  the 
semi-phytophagous  tendencies  of  some  labenines)  that  this  is  partially  a  seed-feeding  ichneumonid  cannot 
be  ruled  out. 

MATERIAL  EXAMINED 

Holotype  $,  Queensland:  Yarraman,  vii.1969  (Heather)  (ANIC). 
1  O"  (paratype),  same  data  as  holotype. 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  139 

POECILOCR  YPTUS  Cameron 

Poecilocryptus  Cameron,  1901:  527.  Type-species:  Poecilocryptus  nigromaculatus  Cameron,  by  mono- 

typy. 
Poecilopimpla  Morley,  1914:  35,  36.  [Unnecessary  replacement  name  for  Poecilocryptus  Cameron.] 

[Homonym  of  Poecilopimpla  Cameron,  1903.] 

Medium-sized  species,  fore  wing  length  6-10  mm;  clypeus  rather  small,  apically  very  thin,  truncate;  labrum 
small,  exposed;  mandible  short,  slightly  twisted,  strongly  narrowed,  bidenjtate;  malar  space  shorter  than 
basal  mandibular  width.  Occipital  carina  dorsally  absent;  eye  with  a  weak  indentation  opposite  antennal 
socket.  Antenna  moderately  long,  clavate.  Mesoscutum  polished,  virtually  impunctate;  notauli  deep  on 
anterior  0-2  of  scutum,  notaular  crests  strong.  Propodeum  abruptly  rounded  with  spiracle  oval;  area 
superomedia  large,  quadrate,  often  confluent  with  area  petiolaris;  area  externa  not  usually  defined 
laterally;  gaster  inserted  well  above  level  of  hind  coxae.  Fore  tibia  simple,  its  apex  not  bearing  a  long  spine; 
tarsal  claws  large,  simple,  or  in  some  species  those  of  the  anterior  two  pairs  of  legs  basally  lobate.  Fore  wing 
with  cu-a  slightly  proximal  to  base  of  Rs&M\  3r-m  complete,  enclosing  a  large,  transverse  pentagonal 
areolet;  Im-cu  with  a  single  bulla,  straight  but  inclivous.  Hind  wing  with  distal  abscissa  of  Cui  absent;  basal 
cell  not  exceptionally  broad;  Sc  bearing  about  two  hamuli.  Gaster  long,  quite  slender;  tergite  1  slender, 
evenly  broadened  posteriorly  with  spiracles  at  or  slightly  behind  centre,  sternite  reaching  to  spiracles,  that 
of  female  bearing  a  pair  of  knob-like  protuberances  near  anterior  end.  Ovipositor  moderately  long, 
projecting  beyond  apex  of  gaster  by  2-8-3-3  times  length  of  hind  tibia,  its  apex  cylindrical,  with  lower  valve 
partially  enclosing  the  upper,  with  an  indistinct  matt  area  laterally,  the  upper  valve  with  weak  dorsal  teeth. 

REMARKS.  Poecilocryptus  is  an  endemic  Australian  genus  with  species  widely  distributed  throughout  the 
continent.  They  seem  to  be  associated  with  a  variety  of  galls  on  trees  of  the  genera  Eucalyptus  and  Acacia. 
Parrott  (1954)  recorded  three  species  from  Australia.  In  this  work  four  species  are  recognized,  two  of 
which  are  described  as  new.  One,  P.  galliphagus,  has  previously  been  incorrectly  known  as  P.  nigromacula- 
tus. In  fact  nigromaculatus  is  a  senior  synonym  of  P.  stramineus. 

The  relationships  of  the  species.  The  four  species  may  be  placed  in  two  species-groups,  the  nigromacula- 
tus-group  (containing  nigromaculatus  and  galliphagus)  which  is  characterized  by  having  acute  lobes  on  the 
claws  of  the  anterior  two  pairs  of  legs,  and  the  nigripectus-group  (containing  nigripectus  and  coloratus) 
which  has  the  first  lateral  area  very  reduced  in  size. 

Key  to  species  of  Poecilocryptus 

1  Tergites  1  and  2  of  gaster  white ;  alitrunk  predominantly  black ,  only  pronotum  and  anterior  parts 

of  mesoscutum  and  mesopleuron  orange coloratus  sp.  n.  (p.  139) 

-  Tergites  1  and  2  of  gaster  bright  yellow,  sometimes  with  black  marks;  alitrunk  predominantly 

bright  yellow  with  profuse  black  spots 

2  Propodeum  with  area  superomedia  delineated  posteriorly  by  a  strong  carina  (Fig.  56) ;  flagellum 

with  a  subapical  pale  mark;  sternite  1  of  gaster  with  weak  antero-ventral  sublateral  keels; 
tergite  2  entirely  yellow nigripectus  Turner  &  Waterston(p.  140) 

-  Propodeum  with  area  superomedia  not  delineated  posteriorly  (Fig.  57);  flagellum  entirely 

black;  sternite  1  of  gaster  with  strong  antero-ventral  sublateral  tubercles;  tergite  2  always 
black-marked 3 

3  Distal  apex  of  hind  femur  black;  ovipositor  projecting  beyond  apex  of  gaster  by  2-0-2-3  times 

length  of  hind  tibia galliphagus  sp.  n.(p.  140) 

-  Distal  apex  of  hind  femur  yellow;  ovipositor  projecting  beyond  apex  of  gaster  by  2-8-3-3  times 

length  of  hind  tibia nigromaculatus  Cameron(p.  141) 

Poecilocryptus  coloratus  sp.  n. 

Female.  Unknown. 

Male.  Small  species,  fore  wing  length  4-5  mm.  Hypostomal  carina,  above  mandibular  base,  weakly 
raised.  Metapleuron  smooth  and  highly  polished.  Propodeal  carinae  quite  weak;  area  superomedia  not 
delineated  by  a  carina  posteriorly;  lateral  longitudinal  carina  present  above  spiracle;  first  lateral  area  quite 
small,  less  than  the  area  of  area  dentipara.  Fore  wing  with  Rs  between  2r-m  and  3r-m  distinctly  shorter  than 
length  of  2r-m.  Fore  and  mid  tarsal  claws  simple.  Gaster  with  sternite  1  simple. 

Coloration.  Face  whitish,  head,  pronotum  and  anterior  parts  of  mesothorax  orange.  Remainder  of 
alitrunk  black.  Gaster  with  tergites  1  and  2  white,  3  white  with  a  central  black  mark,  4+  black  with 
posterior  and  lateral  margins  broadly  white.  Flagellum  black.  Fore  leg  orange;  mid  leg  with  femur  and 


140  I.  D.  GAULD  &  G.  A.  HOLLOW  AY 

proximal  segments  white,  tibia  and  tarsus  blackish;  hind  leg  black,  femur  proximally  and  distally  slightly 
reddish  orange.  Pterostigma  blackish,  wings  infumate. 

REMARKS.  This  small  species  is  easily  recognized  by  its  atypical  colour  pattern.  It  is  the  only  species  in  the 
genus  that  is  not  predominantly  yellow.  The  propodeal  carination  is  also  quite  distinctive.  The  simple  tarsal 
claws  and  small  first  lateral  area  suggest  P.  color atus  may  be  related  to  P.  nigripectus . 

MATERIAL  EXAMINED 

Holotype  cf ,  Tasmania:  Coles  Bay,  ii-iii.19--  (TC). 
Paratypes.  Tasmania:  3  cf ,  same  data  as  holotype  (BMNH,  TC);  1  cf ,  Mt  Barrow,  700  m,  ii.19--  (TC). 

Poecilocryptus  galliphagussp.  n. 

[Poecilocryptus  nigromaculatus  Cameron;  Parrott,  1954:  240.  Misidentification.] 

Female.  Medium-sized  species,  fore  wing  length  7-10  mm.  Hypostomal  carina,  above  mandibular  base, 
moderately  raised.  Metapleuron  smooth  and  highly  polished.  Propodeal  carinae  strong;  area  superomedia 
not  delineated  by  a  carina  posteriorly;  lateral  longitudinal  carina  not  complete  above  spiracle;  first  lateral 
area  very  large,  more  than  twice  the  area  of  area  dentipara.  Fore  wing  with  Rs  between  2r-m  and  3r-m 
about  twice  as  long  as  length  of  2r-m.  Fore  and  mid  tarsal  claws  with  a  well-developed  acute  basal  lobe. 
Gaster  with  sternite  1  bearing  a  pair  of  antero-ventral  sublateral  tubercles.  Ovipositor  projecting  beyond 
apex  of  gaster  by  2-0-2-3  times  length  of  hind  tibia. 

Coloration.  A  bright  yellow  species  with  black  marks  on  interocellar  area,  vertex  behind  ocelli,  three 
longitudinal  stripes  on  mesoscutum,  anterior  propodeal  areae,  anterior  margin  of  all  abdominal  tergites, 
hind  femur  centrally  and  hind  tibia  distally.  Flagellum  and  ovipositor  sheath  black.  Distal  tarsal  segments 
slightly  infuscate.  Pterostigma  dark  brown,  wings  hyaline. 

Male.  Similar  to  female  though  slightly  more  slender,  often  with  sternal  tubercles  very  weak. 

REMARKS.  This  species  is  rather  similar  to  P.  nigromaculatus,  with  which  it  is  frequently  confused  in 
collections.  The  two  species  may  easily  be  separated  by  the  characters  given  in  the  key. 

HOST  RECORDS.  This  species  has  been  reared  from  galls  on  Eucalyptus  delegatensis  and  E.  pauciflora. 

MATERIAL  EXAMINED 

Holotype  $ ,  Victoria:  Wiseleigh  via  Bruthen,  ix.1962,  ex  eucalypt  gall  (Hobb)  (NMV). 

Paratypes.  Queensland:  1  $,  no  further  data  (Riek)  (BMNH).  New  South  Wales:  1  $,  Dainer's  Gap, 
xi.1972  (ANIC);  1  $,  Deer  Vale,  i.1931  (Burns)  (NMV);  1  cf ,  Mt  Victoria,  x.1930  (Burns)  (NMV);  1  $, 
Sydney  (Froggatt)  (ANIC).  Victoria:  2  $,  Eildon  area,  ix.1959  (Irvine)  (NMV);  2  $,  Mt  Pinniber,  iv.1961 
(Taylor)  (ANIC);  1  $,  Toolangi,  xi.1982  (Farrugia)  (BMNH);  3  $,  Warrandyte,  viii-x.1928  (Hill) 
(ANIC);  2  Cf,  no  further  locality  data  (French)  (BMNH).  Tasmania:  3  $,  Collinsvale,  Fairy  Glen, 
i-ii.1983  (Williams  &  Gauld)  (BMNH). 

Poecilocryptus  nigripectus  Turner  &  Waterston 

(Fig.  56) 
Poecilocryptus  nigripectus  Turner  &  Waterston,  1920:  24.  Holotype  $,  TASMANIA  (BMNH)  [examined]. 

Female.  Medium-sized  species,  fore  wing  length  7-11  mm.  Hypostomal  carina,  above  mandibular  base, 
strongly  raised.  Metapleuron  with  distinct  longitudinal  wrinkles,  moderately  polished.  Propodeal  carinae 
strong;  area  superomedia  rectangular,  delineated  by  a  carina  posteriorly;  lateral  longitudinal  carina 
complete  above  spiracle;  first  lateral  area  not  exceptionally  large,  of  approximately  the  same  area  as  area 
dentipara.  Fore  wing  with  Rs  between  2r-m  and  3r-m  subequal  to  length  of  2r-m.  Fore  and  mid  tarsal  claws 
simple.  Gaster  with  sternite  1  bearing  a  pair  of  weak  antero-ventral  sublateral  keels.  Ovipositor  projecting 
beyond  apex  of  gaster  by  2-2-2-4  times  length  of  hind  tibia. 

Coloration.  Bright  yellow  species  with  interocellar  area,  vertex  behind  eyes,  posterior  part  of  mesoscu- 
tum, anterior  part  of  propodeum  and  metapleuron,  anterior  0-7  of  tergite  1,  and  most  of  tergites  3  and  6 
black.  Legs  yellow,  hind  femur  broadly  black  centrally,  distal  hind  tarsal  segment  infuscate.  Flagellum 
black,  with  a  broad  subapical  yellowish  white  band.  Pterostigma  dark  brown,  wings  hyaline. 

Male.  Similar  to  female  but  slightly  more  slender  and  with  gastral  tergites  more  coarsely  punctate. 

REMARKS.  This  is  a  particularly  characteristically  patterned  species.  Structurally  it  is  more  similar  to  P. 
coloratus  than  it  is  to  P.  nigromaculatus,  a  species  that  it  superficially  resembles  in  ground-colour. 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI  141 

HOST  RECORDS.  In  the  BMNH  is  a  specimen  reared  from  anthribid  galls. 

MATERIAL  EXAMINED 

Holotype  $ ,  Tasmania:  Mt  Wellington  (Turner)  (BMNH). 

Queensland:  4  $ ,  Brisbane,  ii.1969  (Campbell)  (QM);  1  cf ,  Iron  Range,  v.1975  (Moulds)  (AM);  1  $,  no 
data  (NMV).  New  South  Wales:  1  cf,  Bateman's  Bay,  x.1969  (Riek)  (ANIC);  1  $>  Bendigo  (Froggatt) 
(ANIC).  Australian  Capital  Territory:  2  $,  Canberra,  x.1930  (Bruce)  (ANIC). 

Poecilocryptus  nigromaculatus  Cameron 

Poedlocryptus  nigromaculatus  Cameron,  1901:  528.  LECTOTYPE  $,  AUSTRALIA  (BMNH),  here  desig- 
nated [examined] . 

Poeciloeryptus  (sic)  nigro-maculatus  Cameron;  Cameron,  1911:  335. 

Poecilopimpla  nigromaculata  (Cameron)  Morley,  1914:  36. 

Poecilopimpla  nigromaculata  var.  straminea  Morley,  1914:  36.  Holotype  $,  NEW  SOUTH  WALES  (BMNH) 
[examined].  Syn.  n. 

Poecilocryptus  stramineus  (Morley)  Parrott,  1954:  241. 

Poecilocryptus  nigromaculatus  Cameron;  Townes  et  al. ,  1961: 117. 

Female.  Medium-sized  species,  fore  wing  length  6-10  mm.  Hypostomal  carina,  above  mandibular  base, 
weakly  raised.  Metapleuron  smooth  and  highly  polished.  Propodeal  carinae  strong;  area  superomedia  not 
delineated  by  a  carina  posteriorly;  lateral  longitudinal  carina  absent  above  spiracle;  first  lateral  area  very 
large,  more  than  twice  the  area  of  area  dentipara.  Fore  wing  with  Rs  between  2r-m  and  3r-m  about  twice  as 
long  as  length  of  2r-m.  Fore  and  mid  tarsal  claws  with  a  well-developed  acute  basal  lobe.  Gaster  with 
sternite  1  bearing  a  pair  of  antero-ventral  sublateral  tubercles.  Ovipositor  projecting  beyond  apex  of  gaster 
by  2-8-3-3  times  length  of  hind  tibia. 

Coloration.  A  bright  yellow  species  with  black  marks  on  interocellar  area,  vertex  behind  ocelli,  three 
longitudinal  stripes  on  mesoscutum,  anterior  propodeal  areae,  anterior  margin  of  all  abdominal  tergites 
and  hind  femur  centrally.  Flagellum  and  ovipositor  sheath  black.  Distal  hind  tarsal  segments  strongly 
infuscate.  Pterostigma  dark  brown,  wings  hyaline. 

Male.  Similar  to  female  though  slightly  more  slender,  often  with  sternal  tubercles  very  weak. 

REMARKS.  In  the  BMNH  are  two  specimens  labelled  as  'Cameron  types'  of  nigromaculatus.  They  are 
conspecific  and  one  has  been  labelled  and  is  here  designated  as  lectotype.  Morley  (1914)  clearly  referred  to 
the  female  that  was  reared  by  Froggatt  as  the  'typical'  specimen,  and  this  must  be  construed  as  a  valid  type 
restriction  (Art.  73(a)(i)  of  the  Code).  Parrott  (1954)  was  incorrect  to  refer  to  the  male  in  Froggatt's 
collection  as  the  holotype.  The  female  holotype  is  a  slightly  undersized  specimen,  but  clearly  conspecific 
with  nigromaculatus. 

P.  nigromaculatus  appears  to  be  the  sister-species  of  P.  galliphagus.  Both  species  have  a  well-developed 
basal  lobe  on  the  fore  and  mid  tarsal  claws  and  have  the  first  lateral  area  of  the  propodeum  greatly 
enlarged.  P.  nigromaculatus  may  be  recognized  by  its  elongate  ovipositor  and  entirely  yellow  hind  tibia. 
We  have  examined  the  differences  between  this  species  and  galliphagus  tabulated  by  Parrott  (1954)  and 
have  found  that  only  the  ovipositor  character  holds  up.  We  failed  to  find  any  difference  in  the  ratio  of 
interocellar  to  orbital-ocellar  distance,  and  the  range  of  numbers  of  flagellar  segments  for  both  species  is 
very  similar.  Small  specimens  of  either  species  have  fewer  flagellar  segments  than  large  individuals.  The 
extent  of  the  black  banding  on  third  to  fifth  gastral  tergites  is  quite  variable  in  both  species. 

HOST  RECORDS.  P.  nigromaculatus  has  been  reared  from  anthribid  and  chalcid  galls  on  Acacia  longifolia, 
and  eriococcid  galls  on  Eucalyptus. 

MATERIAL  EXAMINED 

Lectotype  $  (nigromaculatus  Cameron),  'Australia':  no  further  data  (BMNH).  Paralectotype,  1  $, 
same  data  (BMNH).  Holotype  $  (nigromaculata  var.  straminea  Morley),  New  South  Wales:  no  further 
data  (Froggatt)  (BMNH). 

'Australia':  1  $  (paralectotype  of  nigromaculatus)  (BMNH).  38  $,34  cf,  Queensland,  New  South 
Wales,  Australian  Capital  Territory,  Victoria,  Tasmania  (ANIC,  BMNH,  NMV). 

URANCYLA  Gauld 

Genus  U;  Gauld,  1983:  169. 

Urancyla  Gauld,  1984:  95.  Type-species:  Urancylafulva  Gauld,  by  original  designation. 

Medium-sized  species;  fore  wing  length  6  mm;  clypeus  flat,  small,  apically  truncate  with  margin  thin; 


142  I.  D.  GAULD  &  G.  A.  HOLLO  WAY 

mandible  strongly  tapered,  twisted  and  with  upper  tooth  slightly  the  longer;  malar  space  shorter  than  basal 
mandibular  width.  Occipital  carina  absent  dorsally ,  ventrally  joining  hypostomal  carina  well  above  base  of 
mandible.  Antenna  long,  neither  tapered  nor  clavate  distally.  Mesoscutum  polished,  punctate;  notaulus 
present  near  front  margin,  with  a  small  crest  occluding  extreme  end;  scutellum  flat,  without  lateral  carinae; 
propodeum  evenly  rounded  with  vestiges  of  carinae  though  areae  superomedia  and  petiolaris  are  not 
defined;  gaster  inserted  low  on  propodeum,  near  level  of  hind  coxae.  Fore  tibia  with  a  small  tooth  on  outer 
side,  femur  with  a  weak  longitudinal  ventral  furrow;  tarsal  claws  of  female  with  a  basal  lobe.  Fore  wing  with 
cu-a  subopposite  Rs&M;  3r-m  present,  weakly  pigmented,  enclosing  a  small  pentagonal  areolet;  2m-cu 
with  a  single  bulla.  Hind  wing  with  distal  abscissa  of  Cu\  present;  first  abscissa  of  Cu\  shorter  than  cu-a; 
basal  cell  moderately  broad;  Sc  bearing  one  hamulus.  Gaster  moderately  long,  tergite  1  slender  with 
spiracle  slightly  behind  centre;  tergites  2  and  3  with  pendant  laterotergites  which  are  almost  membranous. 
Ovipositor  about  as  long  as  gaster,  evenly  decurved,  apex  simply  acute  with  inconspicuous  teeth  and 
indistinct  matt  area  laterally. 
Male.  Unknown. 

REMARKS.  In  Townes'  (1969)  key  to  Labiinae  this  genus  runs  to  the  tribe  Clasini  but  it  does  not  appear  to  be 
related  to  the  genera  in  this  group.  Clasines  have  a  nodus  on  the  ovipositor  apex,  long,  simple  claws  and 
two  bullae  in  2m-cu.  They  also  have  no  trace  of  a  notaular  crest.  The  twisted  mandible  and  single  bulla  in 
2m-cu  are  characters  that  Urancyla  shares  with  Poecilocryptus  and  the  two  genera  appear  to  be  closely 
related. 

Urancyla  fulva  Gauld 

(Fig.  64) 
Urancyla  fulva  Gauld,  1984:  95.  Holotype  $,  QUEENSLAND  (TC)  [examined]. 

Female.  Lower  face  elongate,  regularly  punctate;  frons  polished  and  finely  punctate;  ocelli  arranged  in  an 
equilateral  triangle.  Flagellum  with  30  segments.  Mesoscutum  polished,  regularly  and  finely  punctate; 
mesopleuron  highly  polished,  smooth,  almost  impunctate;  metapleuron  polished  with  scattered  fine 
punctures;  submetapleural  carina  moderately  wide,  evenly  tapered  anteriorly.  Gaster  highly  polished, 
finely  punctate. 

Coloration.  Predominantly  orange-brown  species;  face,  upper  orbits,  genae,  propleuron,  anterior 
margin  of  pronotum,  diagonal  stripe  across  mesopleuron,  fore  coxa  and  trochanters  and  a  stripe  on  mid 
coxa  pale  yellowish;  flagellum,  except  centrally,  scape  and  pedicel,  frons  centrally,  vertex,  interocellar 
area,  occiput,  mesoscutum  and  ovipositor  sheath,  pterostigma  black. 

Male.  Unknown. 

REMARKS.  This  species  is  known  only  from  the  holotype. 

MATERIAL  EXAMINED 
Holotype  $,  Queensland:  Brisbane,  xi.1972  (Sedlacek)  (TC). 

Acknowledgements 

We  thank  the  curators  of  the  institutions  that  kindly  loaned  us  material,  and  the  individuals  who  ran 
Malaise  traps  to  secure  additional  specimens.  We  are  particularly  grateful  to  Dennis  Farrugia  for  collecting 
a  considerable  amount  of  material  in  southern  Victoria.  IDG  would  like  to  thank  ABRS  for  providing  a 
grant  to  study  in  Australia.  GAH  would  like  to  thank  the  New  South  Wales  Premier's  Department  for 
support  whilst  overseas,  and  Nestle  (Australia)  Ltd  for  funds. 

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I.  D.  GAULD  &  G.  A.  HOLLOWAY 


14 


Figs  1-14  1,  2,  face  of  (1)  Labium  sp.;  (2)  Labena  sp.  3,  fore  wing  of  Labena  sp.  4,  base  of  ovipositor 
sheath,  Certonotus  nitidulus.  5,  6,  fore  wings  of  (5)  Urancyla  fulva;  (6)  Adelphion  sp.  7,  fore  leg  of 
Labena  sp.  8-10,  gonosquama  of  (8)  Labena  annulata;  (9)  L.  keira;  (10)  L.  pudenda.  11, 12,  hind  wings 
of  (11)  L.  annulata;  (12)  L.  fce/ra.  13, 14,  tergite  2  of  (13)  L.  annulata;  (14)  L.  chadwickii. 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI 


Figs  15-33  15,  16,  end  of  gaster  of  (15)  Labena  grandis;  (16)  L.  malecasta.  17,  18,  mid  tibia  of  (17)  L. 
jacunda;  (18)  L.  keira.  19,  20,  propodeum  of  (19)  L.  keira;  (20)  L.  pudenda.  21,  apex  of  ovipositor,  L. 
pudenda.  22-24,  pro-  and  mesoscutum  of  (22)  Certonotus  rufescens;  (23)  C.  apicalis;  (24)  C.  humeralifer. 
25,  26,  face  of  (25)  C.  rufescens;  (26)  C.  humeralifer.  27-29,  propodeum  of  (27)  C.  humeralifer;  (28)  C. 
annulatus;  (29)  C.  andrewi.  30, 31,  areolet  of  fore  wing  of  (30)  C.  humeralifer,  (31)  C.  annulatus.  32, 33, 
propodeum  of  (32)  C.  geniculatus;  (33)  C.  nitidulus. 


146 


I.  D.  GAULD  &  G.  A.  HOLLOWAY 


Figs  34-49  34,  subalar  prominence  of  Certonotus  talus.  35-37,  end  of  gaster  of  (35)  C.  talus;  (36)  C. 
sisyphus;  (37)  C.  cestus.  38,  39,  mouthparts  and  back  of  head  of  (38)  C.  monticola;  (39)  C.  sisyphus.  40, 
41,  dorsal  view  of  propodeum  of  (40)  C.  sisyphus;  (41)  C.  avitus.  42,  43,  lateral  view  of  propodeum  of 
(42)  C.  avitus;  (43)  C.  cestus.  44,  45,  propodeum  and  tergite  1  of  (44)  C.  celeus;  (45)  C.  pineus.  46,  47, 
alitrunk  of  (46)  C.  pineus;  (47)  C.  mogimbensis.  48,  49,  propodeum  of  (48)  Certonotus  sp.  A;  (49)  C. 
leeuwinensis. 


AUSTRALIAN  LABENINI  &  POECILOCRYPTINI 


147 


Figs  50-64  50,  51,  end  of  gaster  of  (50)  Certonotus  sp.  A;  (51)  C.  zebrus.  52,  53,  head  of  (52)  C. 
leeuwinensis;  (53)  C.  zebrus.  54,  55,  hind  coxa  of  (54)  C.  paluma;  (55)  C.  ixion.  56,  57,  propodeum  of 
(56)  Poecilocryptus  nigripectus;  (57)  P.  nigromaculatus .  58-62,  body  in  profile  to  show  colour  pattern  of 
(58)  Certonotus  farrugiai;  (59)  C.  zebrus;  (60)  C.  toolangi;  (61)  C.  ixion;  (62)  C.  paluma.  63,  fore  wing  of 
Alaothyris  elongissimus .  64,  alitrunk  and  gaster  of  Urancylafulva. 


148 


I.  D.  GAULD  &  G.  A.  HOLLO  WAY 


•H 
0) 


CO 

•8 

0) 


00 


4,5 


10 


7,8 


Fig.  65  Cladogram  showing  putative  phylogenetic  relationship  of  Australian  species  of  Labena.  (Note. 
The  holophyly  of  this  grouping  vis  a  vis  the  Neotropical  species  has  not  been  established.)  The 
apomorphic  features  supporting  this  cladogram  are:  1 ,  mesopleuron  smooth;  2,  ring  of  long  hairs  around 
apex  of  gonosquama;  3,  apex  of  fore  wing  infumate;  4,  apex  of  gonosquama  indented;  5,  apex  of 
ovipositor  with  coarse  teeth;  6,  fore  tibia  slender;  7,  vein  Cu\  incomplete  in  hind  wing;  8,  flagellum  with 
apical  white  band;  9,  gaster  closely  punctate;  10,  mesoscutum  striate. 


Acacia  longifolia  141 
Buprestidae  114, 124 

Cerambycidae  113 
Diadoxus  sp.  124 


INDEX 

Index  to  hosts 


Host  plants,  as  well  as  host  insects,  are  included  in  this  index. 


Ethon  affine  114 
Eucalyptus  141 
Eucalyptus  delegatensis  140 
Eucalyptus  pauciflora  140 

Pinusradiata  114 


Sirex  noctilio  131 
Sirexsp.  114,132 
Siricidae  131 

Ur acanthus  strigosus  113 


149 


Index  to  Ichneumonidae 


Invalid  names  are  in  italics;  principal  references  are  in  bold. 


Alaothyris  109, 138 
andrewi  108, 119, 121 
annulata  109, 112, 113 
apicalis  108, 119, 122 
Asperellus  108, 117 
avitus  108, 120, 123 

celeus  108, 120, 123 
Certonotus  108, 111,117 
cestus  108, 120, 123 
chadwickii  109, 112, 113 
coloratus  109, 139 

elongissimus  109, 138 

farrugiai  108, 120, 126 
fulva  109, 142 

galliphagus  109, 139, 140 
geniculatus  108, 119, 124 
grandis!09, 112, 114 


hinnuleus  108, 120, 126 
humeralifer  108, 119, 127 

ixion  108, 121, 128 
jacunda  109, 112, 115 
keira  109, 112, 115 

Labena  109,  111,  112 
Labeninae  108, 109, 110 
Labenini  108, 109, 110,  111 
leeuwinensis  108, 120, 128 

malecasta  109, 112, 116 
mogimbensis  108, 120, 129 
monticola  108, 119, 130 

nigripectus  109, 139, 140 
nigromaculatus  109, 139, 141 


nitidulus  108, 119, 131 

paluma  108, 121,132 
pineus  108, 120, 132 
Poecilocryptini  108, 109, 137, 138 
Poecilocryptus  109, 138, 139 
pudenda  109, 113, 117 

rufescens  108, 119, 133 

sisyphus  108, 120, 134 
species  A  109, 120, 137 
straminea  109, 141 

talus  108, 119, 121,135 
tasmanienis  108, 131 
toolangi 109, 121, 136 

Uraneyla  109, 138, 141 
zebrus  109, 120, 136 


/.  D.  Gauld 

In  the  important  field  of  biological  and  integrated  control  of  pests  the  parasitic  Hymenoptera 
are  of  particular  significance,  and  this  work  considers  one  of  the  largest  families  of  Parasitica, 
the  Ichneumonidae.  The  group  has  received  little  attention  in  Australia  -  though  it  has  already 
been  utilized  successfully  in  curtailing  the  ravages  caused  by  accidentally  introduced  pests.  For 
selective  control  programmes  to  be  effective,  however,  a  sound  knowledge  of  the  biology  of 
both  the  pest  and  its  parasites  is  essential  -  and  a  sound  taxonomic  base  is  vital  for  the 
development  of  such  knowledge. 

Ironically,  considering  the  group's  economic  importance,  the  parasitic  Hymenoptera  are 
amongst  the  least  studied  of  any  group  of  living  organisms,  and  taxonomic  difficulties  have 
presented  major  problems  to  many  entomologists  working  with  the  Parasitica.  An 
Introduction  to  the  Ichneumonidae  of  Australia  will  go  a  long  way  towards  rectifying  this 
situation,  being  a  taxonomic  treatment,  by  genus,  of  the  Australian  ichneumonids,  a 
comprehensive  illustrated  identification  guide,  and  a  summary  of  all  available  information  on 
the  group.  It  will  also  serve  as  an  introduction  to  the  biology  and  distribution  of  Australian 
ichneumonids,  and  provide  a  check-list  of  the  described  species  and  an  index  to  their  known 
hosts.  It  provides  an  important  revision  of  ichneumonid  nomenclature  in  order  to  bring  the 
group  into  line  with  the  generally  accepted  principles  of  zoological  nomenclature. 

1984, 413pp,  3  maps,  580  figs.  Paperback.  0  565  008%  X  £40.00 


Titles  to  be  published  in  Volume  53 


A  review  of  the  Miletini  (Lepidoptera:  Lycaenidae) 

By  J.N.Eliot 

Australian  ichneumonids  of  the  tribes  Labenini  and  Poecilocryptini 

By  I.  D.  Gauld  &  G.  A.  Holloway 

The  tribe  Pseudophloeini  (Hemiptera:  Coreidae)  in  the  Old  World  tropics  with  a  discussion  on 
the  distribution  of  the  Pseudophloeinae 

By  W.  R.  Dolling 

The  songs  of  the  western  European  grasshoppers  of  the  genus  Omocestusin  relation  to  their 
taxonomy  (Orthoptera:  Acrididae) 

ByD.R.  Ragge 


Photoset  by  Rowland  Phototypesetting  Ltd,  Bury  St  Edmunds,  Suffolk 
Printed  in  Great  Britain  by  Henry  Ling  Ltd,  Dorchester 


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Bulletin  of  the 

British  Museum  (Natural  History) 


The  tribe  Pseudophloeini  (Hemiptera: 
Coreidae)  in  the  Old  World  tropics  with  a 
discussion  on  the  distribution  of  the 
Pseudophloeinae 


W.  R.  Dolling 


Entomology  series 

Vol53  No  3  30  October  1986 


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Entomology  series 
Vol  53  No  3  pp  151-212 


Issued  30  October  1986 


,AL  nia 

The  tribe  Pseudophloeini  (Hemiptera:  Coreidae)  in 
the  Old  World  tropics  with  a  discussion  on  the 
distribution  of  the  Pseudophloeinae 

W.  R.  Dolling 

Department  of  Entomology,  British  Museum  (Natural  History),  Cromwell  Road,  London 
SW7  5BD 

Contents 

Synopsis 151 

Introduction 151 

Diagnosis  of  Pseudophloeinae  and  its  division  into  tribes 152 

Rejected  genera 154 

Abbreviations  of  depositories 155 

Terminology  and  measurements 155 

Key  to  the  genera  of  Pseudophloeini  found  in  the  Old  World  tropics 156 

Risbecocoris  Izzard 156 

HoplolomiaS&l 161 

Indolomia  gen.  n 163 

Paramyla  Linnavuori 165 

Psilolomia  Breddin 168 

Pungragen.n 179 

Pseudomyla  gen.  n 180 

Neomevaniomorpha  gen.  n 183 

Mevaniomorpha  Reuter 185 

Mevanidea  Reuter 187 

Arenocoris  Hahn 191 

A/v/flStal 193 

Distribution  of  Pseudophloeinae 205 

Acknowledgements 209 

References 209 

Index 212 

Synopsis 

The  major  morphological  features  of  the  Pseudophloeini  are  outlined  and  their  bearing  on  the  classifica- 
tion of  the  tribe  is  discussed.  Twelve  genera  and  43  species  are  recognized  in  the  tropical  regions  of  Africa 
and  Asia;  descriptions  and  keys  for  their  separation  are  provided.  Four  genera  and  16  species  are  described 
as  new.  Three  new  synonymies  are  established  at  the  genus-level  and  four  at  the  species-level;  10  new 
combinations  are  established;  two  'forms'  are  raised  to  the  status  of  species  and  nine  lectotypes  are 
designated. 

Introduction 

Plant-feeding  bugs  of  the  family  Coreidae  are  characteristic  inhabitants  of  the  herb  and  shrub 
layers  of  tropical  and,  to  a  lesser  extent,  of  temperate  ecosystems.  They  are  frequently 
encountered  in  surveys  of  crop  pests,  since  most  of  the  world's  tropical  crops  are  herbs  or  shrubs, 
and  are  usually  represented  in  collections  made  during  ecological  studies  in  the  tropics.  Most  of 
the  literature  available  for  the  identification  of  Coreidae  is  out  of  date,  fragmentary  or  lacks 
identification  keys. 

Pseudophloeinae  may  be  recognized,  with  a  little  practice,  by  their  general  habitus  and  size, 
and  by  the  absence  of  a  dorsal  sulcus  on  the  tibiae;  other  groups  of  Coreoidea  lacking  tibial  sulci 


Bull.  Br.  Mus.  not.  Hist.  (Ent.)  53  (3):  151-212  30  October  1986 


152  W.  R.  DOLLING 

-  Alydidae,  Rhopalidae,  Stenocephalidae  and  the  coreid  subfamily  Hydarinae  -  are  unlikely  to 
be  mistaken  for  Pseudophloeinae  because  they  are  all  of  characteristic  appearance.  Coreinae 
similar  in  size  and  build  to  Pseudophloeinae  all  have  very  distinct  tibial  sulci.  Twenty-eight  valid 
genera  and  166  species  of  Pseudophloeinae  have  been  described.  Most  of  the  species  are  7-10 
mm  in  length;  a  species  of  typical  appearance  is  illustrated  in  Fig.  1. 

The  food  plants  of  all  species  of  the  subfamily,  where  known,  are  herbaceous  legumes 
(Fabaceae).  Host  plant  records  for  the  tropical  Pseudophloeini  are  very  scarce,  as  are  other 
details  of  their  biology.  There  is  only  a  single,  New  World,  species  of  the  subfamily  in  which 
facultative  brachyptery  is  known;  all  other  species  are  fully  macropterous.  Flight,  therefore,  is 
probably  important,  yet  there  are  very  few  records  of  them  being  caught  in  light  traps, 
interception  traps  or  yellow  trays;  presumably  they  fly  only  rarely  and  then  not  far.  Most  species 
have  rather  restricted  ranges  and  only  one  is  recorded  from  oceanic  islands.  Tropical-crop 
entomologists  and  collectors  in  Europe  report  them  to  move  sluggishly  even  when  disturbed  and 
to  be  reluctant  to  fly.  The  typical  habitats  of  Pseudophloeinae  reflect  those  of  their  host  plants: 
open  woodlands  and  grasslands  with  scattered  trees;  a  few  species  have  penetrated  drier 
grasslands  on  the  one  hand  and  forest  clearings  on  the  other,  but  deserts  and  dense  forests,  to 
judge  from  the  distribution  patterns  of  the  tropical  species,  are  impenetrable  barriers  to  most  of 
them.  Several  species  are  restricted  to  high  altitudes,  suggesting  that  low  temperatures  are  not 
barriers  to  dispersal  in  the  long  term,  a  reasonable  supposition  in  view  of  the  richness  of  the 
Palaearctic  pseudophloeine  fauna. 

The  subfamily  is  represented  in  almost  all  parts  of  the  major  land  masses  except  for 
non-tropical  Australia.  Recent  revisions  by  Froeschner  (1963),  Dolling  (1977)  and  Dolling  & 
Yonke  (1976)  enable  the  Nearctic  and  Neotropical  species  to  be  identified.  Most  Palaearctic 
species  are  covered  by  the  keys  of  Stichel  (1960),  which  should  be  used  in  conjunction  with  the 
notes  of  Chernova  (1979).  The  largest  Palaearctic  genus,  Coriomeris  Westwood,  was  revised  by 
Chernova  (1978).  A  new  Palaearctic  species  of  Microtelocerus  Reuter  was  described  from  Sinai 
by  Dolling  (19796)  and  a  new  genus  with  a  single  new  species  was  added  by  Puchkov  (1979).  The 
tribe  Clavigrallini,  which  is  confined  to  the  Old  World  tropics  and  contains  a  number  of  pest 
species,  has  been  monographed  by  Dolling  (1978: 1979a).  The  present  revision  covers  all  of  the 
genera  of  Pseudophloeinae  not  treated  in  the  above  works. 

Diagnosis  of  Pseudophloeinae  and  its  division  into  tribes 

Stal's  original  (1868:  535)  diagnosis  of  Pseudophloeinae  mentioned  only  the  presence  of  an 
antevannal  vein.  Later  (1873:  33-34),  he  mentioned  also  the  broad  cells  at  the  base  of  the 
hemelytral  membrane,  the  form  of  the  metathoracic  scent-gland  auricle,  the  lack  of  a  median 
dorsal  impression  on  the  head,  the  prominent  and  gently  declivent  tylus  and  juga,  the  prominent 
posterior  angles  of  the  seventh  abdominal  segment  in  both  sexes,  and  the  non-sulcate  tibiae. 
Dolling  (1978:  282)  gave  a  fuller  diagnosis,  adding  mention  of  the  outer  apical  processes  of  the 
antennifers  and  several  features  of  the  genitalia.  The  antevannal  vein  is  not  present  in  five 
genera:  the  Neotropical  Vilga  Stal;  the  Oriental  Hoplolomia  Stal  and  Indolomia  gen.  n.,  the 
Afrotropical  Paramyla  Linnavuori  and  the  Afro- Asian  Risbecocoris  Izzard.  All  five  of  these 
genera  accord  well  with  the  revised  diagnosis  of  the  subfamily  and  there  seems  no  reason  to 
exclude  them  from  it  because  of  the  absence  of  this  single  character;  in  fact,  Stal  (1873)  included 
Hoplolomia  in  Pseudophloeinae,  presumably  on  the  basis  of  other  features  and  in  ignorance  of 
the  venation  of  the  hind  wing. 

The  tribe  Clavigrallini  was  erected  (as  division  Clavigrallaria)  by  Stal  (1873:  81)  in  a  key  to  the 
African  and  Asian  genera  of  the  subfamily.  It  was  characterized  by  him  as  having  the  scutellum 
convex,  the  base  of  the  posterior  femur  devoid  of  the  small  tubercle  which  is  present  in  many 
genera  of  Pseudophloeinae,  the  body  compressed  laterally,  the  male  genital  capsule  not 
biemarginate  posteriorly,  the  propleuron  emarginate  on  its  posterior  margin  near  the  postero- 
dorsal  angle,  the  second  antennal  segment  equalling  or  exceeding  the  third  in  length  and  the 
prescutellar  angles  of  the  pronotum  armed  with  a  spine.  The  first  couplet  of  Stal's  key  contrasted 
Mevania  Stal,  My  la  Stal  and  Hoplolomia  with  the  'Clavigrallaria',  uniting  these  three  non- 


TRIBE  PSEUDOPHLOEINI 


153 


Fig.  1     Psilolomia  pundaloyae ,  dorsal  view  of  male . 


154  W.  R.   DOLLING 

clavigralline  genera  on  the  basis  of  their  common  possession  of  a  flat  or  almost  flat  scutellum,  a 
tubercle  at  the  base  of  the  posterior  femur,  the  posteriorly  biemarginate  genital  capsule  with  the 
emargination  filled  by  the  apices  of  the  parameres,  the  posterior  tibia  not  or  slightly  shorter  than 
the  femur  and  the  second  antennal  segment  shorter  than  the  third.  Although  Stal  did  not 
formally  diagnose  or  name  a  'division  Pseudophloearia'  to  contrast  with  his  'Clavigrallaria',  his 
recognition  of  the  latter  implies  the  existence  of  the  nominate  tribe.  An  additional  feature,  both 
universal  among  Clavigrallini  and  restricted  to  them,  is  the  presence  of  a  pair  of  small  tubercles 
at  the  base  of  the  mesoscutellum  adjacent  to  the  posterior  margin  of  the  pronotum.  In  a  few 
Clavigrallini,  the  second  antennal  segment  is  shorter  than  the  third,  or  the  scutellum  is  flat,  or 
the  genital  capsule  is  biemarginate  posteriorly,  or  the  posterior  tibiae  are  subequal  in  length  to 
the  femora,  or  the  body  is  depressed  rather  than  compressed.  The  emargination  of  the 
propleuron,  the  absence  of  a  femoral  tubercle  and  the  presence  of  the  basal  tubercles  of  the 
scutellum  are  constant  features  of  the  tribe,  as  is  the  presence  of  an  antevannal  vein.  Among  the 
non-clavigralline  genera,  the  scutellum  is  occasionally  convex  and  the  propleuron  emarginate 
(both  conditions  well  developed  in  Vilga  westwoodi  (Kolenati)).  The  tubercle  at  the  base  of  the 
femur  is  absent  in  Risbecocoris  and  Vilga. 

No  further  tribes  have  been  described,  so  the  subfamily  at  present  comprises  the  Clavigrallini 
and  the  nominate  tribe,  Pseudophloeini.  The  unique  characters  of  the  Clavigrallini  are  probably 
apomorphies  within  Pseudophloeinae  while  those  of  Pseudophloeini  are,  in  general,  plesiomor- 
phies.  Both  the  presence  of  the  basal  tubercle  of  the  posterior  femur  and  the  presence  of  the 
antevannal  vein  of  the  metathoracic  wing  are  probably  apomorphies  within  the  subfamily 
(though  it  is  interesting  to  note  that  the  genus  Spathocera  Stein,  in  the  Coreinae,  has  an 
antevannal  vein,  and  a  group  of  genera  including  Riptortus  Stal,  in  the  Alydinae,  have  a  femoral 
tubercle).  Pseudophloeinae  can,  therefore,  be  divided  into  four  unequal  groups  (Table  1):  Vilga 
and  Risbecocoris,  lacking  both  vein  and  tubercle;  Paramyla,  Hoplolomia  and  Indolomia, 
lacking  the  vein  but  possessing  the  tubercle;  Clavigrallini,  possessing  the  vein  but  lacking  the 
tubercle;  and  all  the  remaining  genera,  possessing  both  vein  and  tubercle.  Assuming  that  the 
lack  of  both  vein  and  tubercle  is  primitive,  a  possible  interpretation  of  the  phylogeny  of  the 
Pseudophloeinae  is  that  Clavigrallini  and  most  Pseudophloeini  are  united  by  descent  from  a 
common  ancestor  that  had  no  femoral  tubercle  but  had  acquired  an  antevannal  vein;  from  this 
stock  developed  two  lines:  on  the  one  hand  Clavigrallini  and  on  the  other  all  those  genera  with 
both  a  femoral  tubercle  and  an  antevannal  vein.  On  this  interpretation,  Paramyla,  Hoplolomia 
and  Indolomia  would  have  acquired  their  femoral  tubercles  independently  of  most  Pseudo- 
phloeini; alternatively,  these  three  genera  may  have  secondarily  lost  their  antevannal  veins. 
Recently,  Stys  (1978)  suggested  that  the  antevannal  vein  was  primitively  present  in  the  common 
ancestor  of  all  Coreidae.  This  interpretation  is  rejected  here  because  it  would  involve  recogni- 
tion of  a  group  comprising  the  genera  Vilga,  Risbecocoris,  Paramyla,  Hoplolomia  and  Indolo- 
mia sharing  the  apomorphy  of  the  loss  of  this  vein.  This  group,  based  on  a  loss  character  that 
unites  these  five  morphologically  diverse  and  geographically  widely  dispersed  genera,  is  unlikely 
to  have  any  phyletic  validity. 

A  classification  that  reflected  the  most  probable  phylogeny  of  the  subfamily  would  require  the 
erection  of  two  or  three  additional  tribes  to  accommodate  Vilga,  Risbecocoris  and,  possibly, 
Paramyla  plus  Hoplolomia  plus  Indolomia.  An  assumption  of  independent  acquisition  of  the 
antevannal  vein  by  Clavigrallini  and  the  main  group  of  Pseudophloeini  would  require  separate 
tribes  for  Vilga  and  Risbecocoris  but  not  the  other  three  of  these  genera.  In  the  present  work 
such  new  tribes  are  not  erected.  The  subfamily  Pseudophloeinae  and  the  tribe  Clavigrallini  are 
both  believed  to  be  holophyletic  in  composition,  leaving  the  nominate  tribe  paraphyletic. 


Rejected  genera 

Four  genera  represented  in  the  geographical  area  covered  by  the  present  revision  were  originally 
described  in  Pseudophloeinae  or  associated  with  genera  that  belong  in  the  subfamily. 

Brotheolus  Bergroth  (1908: 107)  is  a  replacement  name  for  the  preoccupied  Brotheus  Distant 


TRIBE  PSEUDOPHLOEINI  155 

(19020:  248)  which  was  placed  by  Distant  (p.  246)  in  Pseudophloeinae.  It  was  transferred  to 
Coreinae:  Gonocerini  by  Dolling  (19796:  97). 

Trallianus  Distant  (19026:  404-405)  was  originally  placed  in  Pseudophloeinae  but  was 
transferred  to  Coreidae:  Gonocerini  by  Dolling  (19796:  97). 

Cristovallia  Distant  (1920:  149-150)  was  said  by  its  author  to  have  'affinity  with  the  genera 
Clavigralla  and  Ceraleptus'  (both  Pseudophloeinae).  Brown  (1958:  514)  synonymized  its 
type-species,  C.  typica  Distant,  with  Amblypelta  bilineata  Stal  in  Coreinae:  Dasynini. 

Austrocoris  Hsiao  (1965:  426)  was  described  in  Pseudophloeinae.  Later,  Hsiao  (1977:  253) 
synonymized  it  with  Chariesterus  Laporte  but  placed  the  latter  genus  in  Pseudophloeinae.  Its 
correct  position  is  in  Coreinae:  Chariesterini. 

Abbreviations  of  depositories 

Specimens  mentioned  in  the  text  are  held  in  a  number  of  different  depositories;  the  addresses  of 

20  of  these  have  been  abbreviated  as  follows. 

AMNH  American  Museum  of  Natural  History ,  New  York ,  U .  S .  A . 

BMNH  British  Museum  (Natural  History),  London,  U.K. 

BPBM  Bernice  P .  Bishop  Museum ,  Honolulu ,  Hawaii ,  U .  S .  A . 

CAS  California  Academy  of  Sciences ,  San  Francisco ,  U .  S .  A . 

IAR  Institute  of  Agricultural  Research,  Samaru,  Nigeria 

IP  Institut  f iir  Pflanzenschutzforschung ,  Eberswalde ,  D .  D .  R . 

IRSNB  Institut  Royal  des  Sciences  Naturelles,  Brussels,  Belgium 

IZ  Institute  of  Zoology ,  Academy  of  Sciences ,  Leningrad ,  U .  S .  S .  R . 

IZPAN  Institut  Zoologiczny,  Polska  Akademia  Nauk,  Warsaw,  Poland 

MNHN  Museum  National  d'Histoire  Naturelle,  Paris,  France 

MNHU  Museum  fur  Naturkunde  der  Humboldt-Universitat ,  Berlin ,  D .  D .  R . 

MRAC  Musee  Royal  de  1'Afrique  Centrale,  Tervuren,  Belgium 

NMB  National  Museum,  Bulawayo,  Zimbabwe 

NMP  Natal  Museum,  Pietermaritzburg,  South  Africa 

NMV  Naturhistorisches  Museum,  Vienna,  Austria 

NR  Naturhistoriska  Riksmuseet,  Stockholm,  Sweden 

TM  Transvaal  Museum,  Pretoria,  South  Africa 

UG  University  of  Ghana,  Legon,  Ghana 

UM  University  Museum,  Oxford,  U.K. 

ZMU  Zoological  Museum  of  the  University,  Helsinki ,  Finland 

Terminology  and  measurements 

Antennal  and  rostral  segments  are  numbered  I  to  IV  starting  with  the  segment  attached  to  the 
body.  The  term  'rostral'  is  used  in  preference  to  'labial'  because  the  base  of  the  first  segment  of 
the  labium  is  usually  obscured;  the  first  segment  of  the  rostrum  is  arbitrarily  taken  to  commence 
at  the  base  of  the  labrum,  which  is  almost  always  visible.  Measurement  of  antennal  segments 
excludes  the  narrow,  unsculptured  bases  of  segments  I,  II  and  III  and  the  small  ring-segment 
between  HI  and  IV.  Ranges  of  measurements  are  given  where  this  procedure  seems  useful  (it  is 
not  used  in  the  case  of  ratios  of  lengths  of  rostral  and  antennal  segments,  where  it  would  be  too 
cumbersome  and  would  be  of  minimal  use  in  identifying  species).  Means  were  calculated  during 
the  preparation  of  this  revision  but  were  discarded:  because  of  the  unequal  representation  of 
different  populations  in  the  samples  available  it  was  felt  that  they  would  be  unhelpful  and 
possibly  misleading.  Surprisingly  often,  means  were  found  to  fall  almost  exactly  half-way 
between  the  extremes  of  the  ranges  cited.  The  reliability  of  the  range  of  measurements  of  any 
species  as  a  guide  to  what  might  be  encountered  by  the  reader  in  examining  his  own  material  will 
depend  largely  on  the  number  of  specimens  examined  by  the  author  and  the  number  of  localities 
from  which  they  were  collected;  this  information  is  given  under  the  heading  'Material  examined' 
for  each  species.  It  should  be  borne  in  mind  that  the  antennae  of  these  insects  are  fragile  and  that 
the  figures  given  for  the  ratios  of  the  lengths  of  the  antennal  segments  are  in  all  likelihood  based 
on  progressively  fewer  specimens  as  one  proceeds  towards  the  distal  end  of  the  antenna.  The 


156  W.  R.  DOLLING 

term  'posterior'  as  applied  to  angles  and  spines  of  the  pronotum  is  avoided  in  favour  of  the 
unambiguous  terms  'posterolateral'  and  'prescutellar' ;  the  former  usually  project  laterally  or 
anterolaterally  from  the  sides  of  the  body  while  the  latter  are  situated,  if  they  are  present  at  all, 
on  the  posterior  margin  of  the  pronotum  close  to  the  lateral  angles  of  the  scutellum  and  project 
posteriorly.  Detailed  descriptions  of  the  body  sculpture,  pubescence  and  colour  are  usually 
given  either  under  the  description  of  the  genus  or  under  the  description  of  one  species  of  each 
genus  that  typifies  the  condition  of  these  characters  throughout  the  genus,  and  only  deviations 
from  this  pattern,  if  they  occur,  are  given  for  the  other  species.  In  descriptions  of  sculpture,  the 
term  'granule'  is  applied  to  projections  that  are  no  higher  than  their  width  and  'tubercle'  applies 
to  projections  longer  than  this;  on  the  femora,  there  is  a  continuous  gradation  from  granules 
through  tubercles  to  small  spines;  all  of  these  structures  are  probably  derived  from  enlarged  hair 
bases  that,  in  the  case  of  spines,  have  either  lost  the  hair  completely  or  have  its  insertion 
displaced  from  the  apex  of  the  setiferous  tubercle. 

Key  to  the  genera  of  Pseudophloeini  found  in  the  Old  World  tropics 

1  Posterior  femur  without  tubercle  adjacent  to  base  of  trochanter.  Appearance  characteristic 

(Fig.  2) RISBECOCORIS(p.  156) 

-  Posterior  femur  with  tubercle  adjacent  to  base  of  trochanter,  rarely  obsolete  and  insect  then  of 

general  appearance  of  Fig.  1 

2  Antennal  segment  II  less  than  one-third  as  long  as  segment  III ARENOCORIS(p.  191) 

-  Antennal  segment  II  more  than  half  as  long  as  segment  III 3 

3  Posterior  margin  of  pronotum  with  a  pair  of  spines  projecting  backwards  over  bases  of  clavi  at 

rest  (Figs  20, 25, 77) 4 

-  Posterior  margin  of  pronotum  smooth  or  at  most  with  a  few ,  low  tubercles  or  granules 5 

4  Head  dorsally  and  pronotum  laterally  with  long  spines;  antennal  segment  I  strongly  clavate, 

with  many  long  spines  and  tubercles  (Fig.  77) MEVANIDEA(p.  187) 

-  Head,  pronotum  and  antennae  without  spines PARAMYLA (p.  165) 

5  Main  pubescence  of  body  and  hemelytra  of  short ,  decumbent ,  scale-like  hairs  (Figs  97 , 98) 6 

-  Main  pubescence  of  body  and  hemelytra  of  longer ,  erect  or  suberect ,  bristle-like  hairs 8 

6  Male  with  apical  tooth  of  paramere  upcurved,  apex  of  paramere  not  filling  posterior  emargina- 

tion  of  genital  capsule  (Fig.  96).  (Africa) MYLA(p.  193) 

-  Male  with  parameres  club-shaped,  apical  tooth  short  and  not  curved,  apex  of  paramere  filling 

posterior  emargination  of  genital  capsule  (as  in  Fig.  21) .  (Asia) 7 

7  Pronotum  with  posterolateral  angles  strongly  produced  anterolaterally  (Figs  63,  64). 

PSEUDOMYLA(p.  180) 

-  Pronotum  with  posterolateral  angles  slightly  prominent  (Fig.  58) PUNGRA  (p.  179) 

8  Abdominal  sternites  III  to  VII  with  posterolateral  angles  right-angled  or  acute,  not  projecting 

as  triangular  teeth  (maximum  degree  of  serration  as  in  Fig.  42) PSILOLOMIA  (p.  168) 

-  Abdominal  sternites  III  to  VII  produced  into  triangular  teeth,  making  outline  of  abdomen 

coarsely  serrate  (Figs  13,19) 

9  Antevannal  vein  present  in  metathoracic  wing.  (Africa) 10 

-  Antevannal  vein  absent  from  metathoracic  wing.  (Asia) 11 

10  Scutellum  terminating  apically  in  a  small,  elevated,  white  blob MEVANIOMORPHA  (p.  185) 

-  Scutellar  apex  pointed,  neither  elevated  nor  white NEOMEVANIOMORPHA  (p.  183) 

11  Pronotum  (Fig.  12)  coarsely  granulate-tuberculate,  posterolateral  spines  arising  abruptly  from 

posterolateral  angles HOPLOLOMIA(p.  161) 

-  Pronotum  (Fig.  18)  finely  granulate-tuberculate,  posterolateral  angles  tapering  smoothly  into 

posterolateral  spines INDOLOMIA(p.  163) 

RISBECOCORIS  Izzard 

Risbecocoris  Izzard,  1949: 478-479.  Type-species:  Risbecocoris  tomentosus  Izzard,  by  original  designation. 

Body  rather  elongate,  about  2-5-3-0  times  as  long  as  broad,  strongly  depressed.  Connexivum  moderately 
expanded. 

Head  about  as  long  as  pronotum,  strongly  convex;  eyes  small,  prominent;  ocelli  dorsally  obscured  by 
pads  of  tomentose  pubescence;  dorsum  of  head  strongly  granulate  and,  at  level  of  base  of  antennifers,  with 
a  pair  of  prominent  setiferous  tubercles;  each  antennifer  laterally  with  two  or  three  similar  tubercles  and  a 


TRIBE  PSEUDOPHLOEINI  157 

broad,  weakly  deflexed  and  ventrally  incurved  apical  process.  Antennal  segment  I  varying  in  length  from 
about  four- fifths  to  almost  equal  to  head  width  including  eyes,  with  long,  outstanding  tubercles,  cylindrical 
through  most  of  its  length  but  narrowed  gradually  towards  base  in  proximal  one-quarter;  segments  II  and 
III  distinctly  more  slender  than  I,  tuberculate,  II  shorter  than  I,  III  longest  of  all;  IV  shortest,  slightly 
thicker  than  II  or  III,  elongate  fusiform,  specialized  sensory  area  occupying  its  apical  two-thirds.  Bucculae 
occupying  about  one-half  of  ventral  midline  of  head.  Rostrum  at  rest  reaching  to  posterior  margin  of 
mesosternum,  segments  I  and  II  subequal,  HI  about  half  as  long  as  I  and  IV  about  two-thirds  as  long  as  I. 
Posterior  half  of  head  with  two  rows  of  tubercles  flanking  rostrum,  in  line  with  bucculae. 

Pronotum  shallowly  declivent,  granulate,  posterior  margin  straight,  prescutellar  spines  absent,  postero- 
lateral  angles  weakly  elevated,  lateral  margins  almost  straight,  with  five  long,  laterally  directed  spines. 
Scutellum  about  1-2  times  as  long  as  its  basal  width,  almost  flat,  apex  pointed.  Mesosternum  deeply  sulcate 
longitudinally.  Metasternum  convex  with  fine,  median,  longitudinal  groove.  Metathoracic  scent-gland 
peritreme  with  dorsal  ridge  modified  into  a  short,  spout-like  structure,  circular  in  outline  and  completely 
surrounding  orifice  (Fig.  4).  Corium  with  costal  margin  convex,  apical  margin  weakly  convex,  all  veins  very 
strongly  prominent;  membrane  of  hemelytron  with  venation  reticulate,  prominent.  Metathoracic  wing 
(Fig.  5)  with  subcosta  free  in  apical  two-thirds,  antevannal  vein  absent.  Femora  and  tibiae  with  rows  of 
prominent  tubercles  or  granules;  posterior  femur  without  subapical  spines  but  with  one  or  two  small 
tubercles  in  this  position  and  no  apical  series;  base  of  femur  adjacent  to  trochanter  lacking  a  tubercle. 

Abdominal  sternites  III-VII  with  posterolateral  angles  produced  into  progressively  longer  spines, 
lateral  margins  of  sternites  and  spines  bearing  prominent  granules  and  tubercles.  Abdominal  spiracles 
situated  very  close  to  lateral  margins  of  sternites,  prominent.  Male  genital  capsule  (Figs  9-11)  broadly 
emarginate  posteriorly,  emargination  filled  by  apices  of  parameres.  Phallotheca  with  a  ventral  sclerite, 
produced  dorsolaterally,  and  two  rather  broad  dorsal  longitudinal  sclerites.  Conjunctive  with  median 
dorsal  and  ventral  lobes,  distal  and  apical  lobes  various.  Vesica  short,  not  protected  by  any  basal  sclerites, 
wings  of  ejaculatory  reservoir  complex  articulating  with  broad,  distal  dorsolateral  sclerites.  Spermathecal 
duct  short,  almost  straight  except  for  one  sharp  bend;  bulb  narrowly  lunate.  Anus  of  female  directed 
ventrally. 

Dense,  off-white,  tomentose  pubescence  present  on  head,  pronotum,  scutellum,  clavus,  corium, 
connexivum,  underside,  antennae  up  to  and  including  base  of  segment  IV  and  legs  up  to  and  including 
bases  of  tibiae.  Erect  hairs  of  body  and  of  first  one  or  two  antennal  segments  very  long  and  curved, 
remaining  parts  of  antennae  and  other  appendages  with  shorter,  straight,  erect  hairs. 

REMARKS.  All  species  of  this  genus  have  a  very  distinctive  appearance  (Fig.  2)  by  virtue  of  their  long,  dense 
pubescence  and  long,  lateral  pronotal  spines.  Reticulate  venation  of  the  forewing  membrane  is  characteris- 
tic of  ground-dwelling  Coreidae.  The  distally  free  subcosta  in  the  metathoracic  wing  is  unique  in  the  family. 
The  features  that  give  the  insects  of  this  genus  their  remarkable  appearance  have  probably  arisen  in 
response  to  the  demands  of  what  is,  for  a  coreid,  an  unusually  arid  habitat. 

DISTRIBUTION.  All  records  come  from  a  belt  of  semi-arid  terrain  stretching  across  Africa  from  Senegal  to 
Kenya  and  thence  into  the  Indian  Desert. 

Key  to  species 

1  Larger,  length  of  male  7-2  mm  or  more,  of  female  7-7  mm  or  more;  antennal  segment  III  about 

1-3  times  as  long  as  segment  I.  (Sudan,  Nigeria,  Kenya,  Chad) numidianus(p.  160) 

-  Smaller,  length  of  male  7-1  mm  or  less,  of  female  7-6  mm  or  less;  antennal  segment  III  between 

1  •  1  and  1-2  times  as  long  as  segment  1 2 

2  Posterior  femur  with  low  granules  on  dorsal  surface.  (Niger) airensis(p.  159) 

-  Posterior  femur  with  outstanding  tubercles  on  dorsal  surf  ace 3 

3  Spines  of  posterolateral  angles  of  abdominal  segments  with  weak  sigmoid  curvature,  those  of 

segment  VII  apically  divergent.  (Pakistan) quadrocephalus(p.  161) 

-  Spines  of  posterolateral  angles  of  abdominal  segments  with  stronger  sigmoid  curvature ,  those  of 

segment  VII  apically  convergent 4 

4  Larger,lengthofmale6-7-7-lmm,offemale7-l-7-6mm. (Senegal) tomentosus(p.  157) 

-  Smaller,  length  of  female  6-4-6-7  mm,  male  unknown.  (India) delhiensis(p.  160) 

Risbecocoris  tomentosus  Izzard 

(Figs  5-11) 
Risbecocoris  tomentosus  Izzard,  1949:  479-480,  pi.  7.  Holotype  cf ,  SENEGAL  (BMNH)  [examined]. 


158 


W.  R.  DOLLING 


Figs  2-11  Risbecocoris  species.  2,  3,  numidianus:  (2)  dorsal  view;  (3)  apical  view  of  conjunctiva  and 
vesica.  4,  quadrocephalus ,  left  metapleuron  and  surrounding  area,  showing  scent-gland  peritreme.  5-11, 
tomentosus:  (5)  metathoracicwing;  (6)  dorsal  view  of  conjunctiva  and  vesica;  (7)  apical  view  of  same;  (8) 
ventral  view  of  same;  (9)  posterior  view  of  male  genital  capsule  with  parameres;  (10)  dorsal  view  of 
same;  (11)  lateral  view  of  same. 


TRIBE  PSEUDOPHLOEINI  159 

Length:  <D",  6-7-7-1  mm;  $,7-1-7-6 mm. 

Head  quadrate,  its  dorsal  spine-like  tubercle  shorter  than  diameter  of  eye;  tubercles  on  lateral  margin  of 
antennifer  about  4  in  number,  conspicuous,  2-4  times  as  long  as  wide.  Antenna  with  first  segment  about 
0-95  times  as  long  as  width  of  head  including  eyes;  ratio  of  lengths  of  segments  about  1-00:0-93: 1-11:0-56. 
Tubercles  of  segment  I  about  twice  as  long  as  wide,  those  of  segment  II  about  1-5  times  as  long  as  wide. 
Ratio  of  lengths  of  rostral  segments  about  l-00:0-90:0-43:0-67.  Apex  of  corium  at  rest  reaching  to  basal 
quarter  or  basal  third  of  laterotergite  VI.  Width  across  hemelytra  at  rest  about  0-72  times  abdominal  width 
(excluding  spines  and  tubercles)  in  male,  0-77  times  in  female;  abdominal  width  about  0-38  times  total  body 
length.  Posterolateral  spines  of  abdominal  sternites  with  strong  sigmoid  curvature,  those  of  segment  VII 
apically  slightly  convergent.  Tubercles  on  margins  of  abdominal  sternites  1-50-2-25  times  as  long  as  wide, 
absent  from  segments  I-IV,  absent  or  single  on  segment  V,  one  or  two  (rarely  absent)  on  segment  VI,  one 
or  two  on  segment  VII.  All  femora  dorsally  with  two  rows  of  short  tubercles. 

Conjunctiva  (Figs  6-8)  with  low,  M-shaped  dorsomedian  lobe;  large,  membranous,  paired  distal 
dorsolateral  lobes;  globose,  membranous  apical  dorsal  lobes,  small,  membranous,  digitiform  apical 
ventral  lobes,  globose,  membranous  distal  ventrolateral  lobes  and  a  small,  median,  apically  bifid 
ventromedian  lobe.  Distal  ventrolateral  lobes  each  supported  by  a  slender  sclerite  along  its  distal  wall,  this 
sclerite  projecting  anteriad  as  a  narrowly  triangular  appendage,  probably  representing  the  totally 
sclerotized  mesal  lobe  of  the  bilobed  distal  ventrolateral  lobe.  Distal  dorsolateral  lobes  each  with  a  strong, 
heavily  sclerotized  appendage  arising  near  its  base,  adjacent  to  the  apex  of  the  respective  wing  of  the 
ejaculatory  reservoir  complex,  these  sclerotized  appendages  not  expanded  apically. 

Overall  coloration  appearing  pale  grey-brown  to  the  naked  eye.  Colour  of  integument  mid-brown; 
corium  between  veins,  tibiae  except  for  basal  and  apical  annuli  and  spines  of  head  yellow;  lateral  spines  of 
pronotum  yellow  with  apices  black;  apices  of  abdominal  spines  black.  Forewing  membrane  slightly  milky 
hyaline,  veins  white  with  occasional  brown  markings. 

REMARKS.  This  species  differs  from  R.  numidianus  in  its  smaller  body  size  and  less  elongated  antennal 
segments  III  and  IV,  as  well  as  in  the  form  of  the  sclerotized  distal  appendages  of  the  conjunctiva;  the  other 
known  African  species,  R.  airensis,  has  the  tubercles  of  the  femora  reduced  to  granules  only  about  as  high 
as  wide  on  the  anterior  two  pairs  of  legs  and  obsolete  on  the  posterior  pair,  whereas  in  tomentosus  these 
tubercles  are  distinctly  longer  than  wide. 

DISTRIBUTION.  Known  only  from  the  type-locality  in  Senegal. 

MATERIAL  EXAMINED 

Senegal:  1  C?  (holotype),  Bambey,  26.ix.1942  (/.  Risbec)  (BMNH). 

Senegal:  5  C?,  7  $  (paratypes),  Bambey,  26.ix.1942  [Izzard,  1949,  gives  dates  as  ix.1940, 26.xi.1943  and 
1946.]  (BMNH). 


Risbecocoris  airensis  Villiers 
Risbecocoris  airensis  Villiers,  1950a:  323-324.  Holotype  $ ,  NIGER  (MNHN)  [examined]. 

Length:  cf,  unknown;  9 ,  7-2  mm. 

Very  similar  to  R.  tomentosus.  First  antennal  segment  as  long  as  head.  Ratio  of  lengths  of  antennal 
segments  as  1-00:0-96: 1-18:0-68.  Tubercles  on  lateral  margin  of  antennifer  short,  hidden  among  the 
pubescence.  Dorsal  spine-like  tubercle  of  head  much  shorter  than  diameter  of  eye.  Lateral  spines  of 
holotype  all  with  apices  broken  (not  short  as  stated  in  original  description).  Anterior  and  intermediate 
femora  with  two  dorsal  rows  of  hispid  granules  about  as  high  as  wide;  posterior  femora  with  granules 
obsolete.  Apex  of  corium  at  rest  just  reaching  base  of  laterotergite  VI.  Lateral  margins  of  abdominal 
sternites  I-V  without  long  tubercles,  VI  with  a  tubercle  on  one  side  only,  VII  with  a  tubercle  on  both  sides. 
Width  of  abdomen  (excluding  spines)  0-38  times  body  length;  width  across  closed  hemelytra  0-82  times 
abdominal  width. 

REMARKS.  This  species  can  be  distinguished  from  others  of  the  genus  by  the  obsolete  granulation  of  the 
posterior  femora.  The  male  is  unknown;  the  holotype,  stated  to  be  of  this  sex  in  the  original  description,  is 
in  fact  a  female  and  it  remains  the  only  known  specimen. 

DISTRIBUTION.  Air  Mountains. 

MATERIAL  EXAMINED 
Niger:  1  $  (holotype),  Air  Sud,  Agadez,  525  m,  l-5.viii.1947  (A.  Villiers  &  L.  Chopard)  (MNHN). 


160  W.  R.  DOLLING 

Risbecocorisnumidianussp.  n. 

(Figs  2,  3) 

Length:  cf, 7-2-8-0 mm;  $,7-7-8-3 mm. 

Head  quadrate,  dorsal  spine-like  tubercles  longer  than  diameter  of  eye;  antennifers  laterally  with  4-6 
conspicuous  tubercles  up  to  5  times  as  long  as  wide.  Antennae  with  segment  I  0-89-0-95  times  as  long  as 
width  of  head  including  eyes;  ratio  of  lengths  of  segments  about  1-00:0-90: 1-34:0-72.  Tubercles  of  antennal 
segment  I  twice  as  long  as  wide  or  more,  those  of  II  up  to  twice  as  long  as  wide,  those  of  III  and  base  of  IV 
about  as  long  as  wide.  Ratio  of  lengths  of  rostral  segments  about  l-00:0-90:0-43:0-52. 

All  femora  dorsally  with  two  rows  of  tubercles  about  2-5  times  as  long  as  wide,  tubercles  of  tibiae  about 
1-5  times  as  long  as  wide.  Apex  of  corium  at  rest  reaching  one-third  or  one-half  of  the  way  along 
laterotergite  VI.  Spines  of  posterolateral  angles  of  abdominal  segments  with  weak  sigmoid  curvature, 
apices  of  spines  of  segment  VII  weakly  divergent.  Numbers  of  tubercles  present  on  lateral  margins  of 
abdominal  sternites  (excluding  those  on  the  spines  themselves)  varying  from  none  to  three  on  segments  III 
and  IV,  one  to  three  on  V  and  VI  and  three  to  five  on  VII. 

Conjunctiva  (Fig.  3)  differing  from  that  of  tomentosus  in  that  apical  dorsal  and  apical  ventral  lobes  are 
absent  and  sclerotized  appendages  of  distal  dorsolateral  lobes  are  broadly  expanded  apically. 

Overall  coloration  dark  grey-brown;  integument  generally  dark  brown;  forewing  membrane  milky 
hyaline,  its  veins  white  with  some  brown  streaks;  tibiae,  except  for  basal  and  apical  annuli,  yellow-brown; 
corium  and  clavus  between  veins  pale  brown;  tubercles  of  head,  abdomen  and  appendages  pale  brown; 
lateral  spines  of  pronotum  and  abdomen  pale  brown,  black-tipped;  antennal  segment  IV  very  dark 
red-brown. 

REMARKS.  This  is  the  largest  species  of  the  genus.  It  differs  from  all  its  relatives  in  the  proportionately 
longer  antennal  segments  III  and  IV  and  in  the  apically  expanded  distal  sclerotized  appendages  of  the 
conjunctiva;  a  distinctive  species  in  an  otherwise  structurally  rather  uniform  genus. 

DISTRIBUTION.  Widespread  in  the  eastern  Sahel. 

MATERIAL  EXAMINED 

Holotype  cf ,  Sudan:  Umm  Berembeita  [H°51'  N,  30°40'  E;  spelt  Umberumbeita  on  label],  on  ground, 
22.iii.1931  (F.  G.  S.  Whitehead)  (BMNH). 

Paratypes.  Sudan:  1  $,  Upper  Nile  Province,  Steamer  Falweil,  swamps,  21.x. 1933  (A.  D.  Sherati) 
(BMNH);  2  $,  U.N.P.,  Renk-Malakal,  2-^.i.l963  (R.  Linnavuori)  (BMNH  and  R.  Linnavuori  coll.);  1 
Cf ,  1  $,  U.N.P.,  nearMalakal,  5-20.U963  (R.  Linnavuori);  1  cf ,  Blue  Nile,  IngessanaMts,  17-22. id.  1962 
(R.  Linnavuori);  1  cf,  Blue  Nile,  Umm  Banein,  14. xi. 1962  (R.  Linnavuori)  (all  R.  Linnavuori  coll.). 
Chad:  1  cf ,  Bas-Chari,  near  Fort-Lamy,  Farcha,  Forest,  8.viii.l963  (J.  Pericart);  1  cf ,  Bas-Chari,  near 
Douggia,  13.viii.1963  (/.  Pericart)  (both  R.  Linnavuori  coll.).  Nigeria:  1  $,  NW.  State,  Badeggi  RRS, 
19.iii.1972  (/.  T.  Medler)  (University  of  Ibadan).  Kenya:  1  <j>,  Northern  Frontier  District,  Wajeir, 
24.U955,  at  light  (/.  Lansbury)  (BMNH). 

Risbecocoris  delhiensis  (Bose)  comb.  n. 

Hoplolomia  delhiensis  Bose,  1946:  75-76.  Holotype  (sex  unknown),  INDIA:  Delhi,  ix.1937,  at  light  (Kerr) 
(Institute  of  Agricultural  Research,  New  Delhi)  [not  examined]. 

Length:  cf ,  unknown;  $ ,  6-4-6-7  mm. 

Very  similar  to  R.  tomentosus.  Dorsal  spine  of  head  much  shorter  than  diameter  of  eye;  antennifers 
bearing  four  tubercles  up  to  3  times  as  long  as  wide  on  their  lateral  margins.  Head  slightly  wider  than  long. 
First  segment  of  antennae  distinctly  shorter  than  head,  about  0-85  times  as  long  as  head  width  including 
eyes;  ratio  of  lengths  of  antennal  segments  about  1-00:0-85:1-14:0-71.  Tubercles  of  antennal  segment  I  at 
most  twice  as  long  as  wide,  of  segment  II  about  1-5  times  as  long  as  wide,  of  segment  III  about  as  long  as 
wide.  Ratio  of  lengths  of  rostral  segments  about  1-00:0-87:0-54:0-71.  Femora  with  two  dorsal  rows  of 
tubercles  about  1-5  times  as  long  as  wide,  tibiae  with  granules  about  as  high  as  wide.  Abdominal  sternites 
with  posterolateral  spines  stout  and  with  rather  strong  sigmoid  curvature,  those  of  segment  VII  slightly 
convergent.  Abdominal  margins  between  the  spines  (which  themselves  bear  tubercles)  with  tubercles  up  to 
3  times  as  long  as  wide,  borne  singly  on  segments  V-VII.  Male  unknown. 

REMARKS.  In  the  original  description,  the  body  length  is  given  as  7  mm.  This  species  is  the  smallest  of  the 
genus.  The  holotype,  said  to  be  in  the  Agricultural  Research  Institute  at  New  Delhi,  was  not  available  for 
study;  the  paratype,  allegedly  deposited  in  the  BMNH,  was  not  found. 


TRIBE  PSEUDOPHLOEINI  161 

DISTRIBUTION.  Eastern  fringes  of  the  Indian  Desert. 

MATERIAL  EXAMINED 
India:  2  $,  Rajasthan,  Pilani,  ix.1965  (5.  C.  Goel)  (BMNH). 

Risbecocoris  quadrocephalus  Ahmad  &  Shadab 
(Fig.  4) 

Risbecocoris  quadrocephalus  Ahmad  &  Shadab,  1969:  151-155.  Holotype  $,  PAKISTAN:  Karachi,  Malir, 
on  grass  near  lucerne  field,  7.12.1967  (Fareed  Ahmed)  (Karachi  University  Natural  History  Museum) 
[not  examined]. 

Length:  cT,  6-7  mm;  $,7-2-7-3  mm. 

Similar  to  R.  delhiensis  but  slightly  larger  and  tubercles  of  body  and  appendages  slightly  longer.  Length 
of  antennal  segment  I  divided  by  head  width  including  eyes  0-88-0-92.  Ratio  of  lengths  of  antennal 
segments  I:II:III  about  1-00:0-89:1-17;  length  of  segment  IV  divided  by  that  of  segment  I  0-66  (type)  or 
0-69  in  female,  0-77  in  male.  Posterolateral  spines  of  abdominal  sternite  VII  slightly  divergent  apically.  A 
full  description  of  the  female  is  given  in  the  original  description.  Conjunctiva  of  male  similar  to  that  of  R. 
tomentosus  in  all  respects  except  that  the  membranous  lobe  of  the  distal  ventrolateral  lobes  is  smaller  and 
the  ventromedian  lobe  is  somewhat  larger. 

REMARKS.  The  differences  between  the  Asian  species  of  Risbecocoris  are  very  slight.  Substantially  more 
material  would  be  required  for  a  definitive  treatment  of  these  forms.  The  orientation  of  the  spines  of 
abdominal  sternite  VII  can  be  seen  in  the  figures  given  in  the  original  descriptions  of  delhiensis  and 
quadrocephalus  and  may  be  reliable  characters  for  distinguishing  these  two  species.  The  two  described 
Asian  species  are  structurally  very  similar  to  tomentosus  and  the  Arabian  specimen  mentioned  below. 

DISTRIBUTION.  Southern  Pakistan. 

MATERIAL  EXAMINED 

Pakistan:  1  d",  1  $,  Sind,  Mainiforest,  grass,  7.xi.l975  (Ali  Khan)  [determined  as  quadrocephalus  by 
Ahmad  and  Khan]  (BMNH). 

Risbecocoris  sp. 

A  single  damaged  specimen  is  mentioned  here  because  of  its  biogeographical  interest.  It  is  a  male  from  the 
island  of  Kamaran  in  the  People's  Democratic  Republic  of  the  Yemen  ('South  Yemen'),  collected  by  G.  C. 
Champion  and  deposited  in  the  BMNH.  It  is  6-5  mm  long;  the  length  of  its  first  antennal  segment  is  0-90 
times  the  width  of  the  head  including  the  eyes  and  the  second  antennal  segment  is  0-87  times  as  long  as  the 
first;  the  remaining  antennal  segments  are  missing.  Structurally,  it  is  identical  in  almost  every  detail  to  R. 
tomentosus,  including  the  detailed  structure  of  the  conjunctiva.  The  only  differences  apparent  are  the  more 
slender  vesica,  the  slightly  smaller  size  and  the  slightly  paler  colour.  The  presence  of  a  tomentosus-like  form 
on  an  island  near  the  Arabian  shore  of  the  Red  Sea  suggests  that  similar  forms  were  and  perhaps  still  are 
present  throughout  the  northern  edge  of  the  Afrotropical  Region  and  it  forms  a  link  between  the 
Senegalese  and  Indo-Pakistan  species. 

HOPLOLOMIA  Stal 
Hoplolomia  Stal,  1873:  82,  84.  Type-species:  Hoplolomia  scabricula  Stal,  by  monotypy. 

Body  oblong,  slightly  depressed,  connexivum  greatly  expanded,  insect  strongly  spinose-tuberculate  and 
hispid. 

Head  slightly  longer  than  pronotum,  tuberculate;  eyes  rather  small.  Antennifers  strongly  divergent, 
their  outer  apical  processes  directed  obliquely  forwards  and  downwards,  slightly  curved  inwards  apically. 
Antennal  segment  I  clavate,  tuberculate;  segment  III  the  longest,  IV  shortest,  fusiform,  specialized 
sensory  area  of  IV  occupying  three-quarters  of  its  length.  Bucculae  rather  short,  occupying  somewhat  less 
than  one-third  of  ventral  midline  of  head.  Rostrum  at  rest  reaching  disc  of  metasternum;  segments  I  and  II 
subequal,  IV  shorter,  III  very  short. 

Pronotum  (Fig.  12)  moderately  or  weakly  declivent,  tuberculate;  posterolateral  angles  somewhat 
produced,  bearing  spines;  posterior  margin  almost  straight,  prescutellar  spines  absent.  Scutellum 
equilateral,  flat  or  weakly  convex,  apex  slightly  raised.  Dorsal  ridge  of  metathoracic  peritreme  reniform, 
prominent.  Meso-  and  metasternum  shallowly  sulcate.  Corium  with  costal  margin  shallowly  concave, 


162  W.  R.  DOLLING 

apical  margin  straight,  apex  not  produced.  Metathoracic  wing  (Fig.  17)  without  antevannal  vein.  Femora 
and  tibiae  strongly  granulate  to  tuberculate;  posterior  femur  with  small  basal  tubercle,  distally  (Fig.  14) 
with  two  major  spines  and  an  apical  series  of  three  or  four. 

Abdominal  sternites  with  lateral  margins  tuberculate,  posterolateral  angles  spinously  produced  (Fig. 
13).  Male  genital  capsule  posteriorly  emarginate,  emargination  filled  by  apices  of  parameres.  Conjunctiva 
(Figs  15,  16)  with  ventral  lobes  denticulate.  Spermatheca  with  bulb  narrowly  lunate,  duct  about  twice  as 
long  as  bulb,  simply  looped. 

REMARKS.  This  small  genus  is  characterized  by  the  presence  of  a  basal  tubercle  on  the  posterior  femur  in 
conjunction  with  the  absence  of  an  antevannal  vein  in  the  metathoracic  wing  and  the  strongly  tuberculate 
and  hispid  body.  Hoplolomia  delhiensis  Bose  is  transferred  to  Risbecocoris  (p.  160). 

DISTRIBUTION.  Oriental  region. 

Key  to  species 

1     Scutellum  flat.  (India) campbelli(p.  163) 

Scutellum  distinctly  convex.  (South  East  Asia) scabricula(p.  162) 

Hoplolomia  scabricula  Stal 

(Figs  12-14) 
Hoplolomia  scabricula  Stal,  1873:  84.  Holotype  $,  'INDIA'  (NR)  [examined]. 

Length:  cf,  $,6-4 mm. 

Length  of  antennal  segment  I  about  0-93  times  as  long  as  width  of  head  including  eyes ;  ratio  of  lengths  of 
antennal  segments  about  l-00:0-81:l-20:0-62  (female).  Ratio  of  lengths  of  rostral  segments  about 
l-00:0-96:0-38:0-67. 

Pronotum  (Fig.  12)  moderately  declivent,  spines  of  posterolateral  angles  directed  laterad  and  slightly 
anteriad,  width  across  apices  of  spines  divided  by  width  of  head  including  eyes  2-17-2-54.  Scutellum  weakly 
but  distinctly  convex. 

Conjunctiva  with  dorsomedian  lobe  deeply  cleft,  appearing  almost  as  a  pair  of  membranous  dorsolateral 
lobes;  distal  dorsomedian  lobe  small,  conical,  membranous;  distal  dorsolateral  lobes  rather  globular,  each 
with  a  finger-like,  membranous  lobe  arising  at  junction  with  dorsomedian  lobe  and  a  short,  curved, 
sclerotized  appendage  arising  opposite  apex  of  wing  of  ejaculatory  reservoir  apparatus  and  curving 
downwards;  apical  ventral  lobes  membranous,  globular;  distal  ventrolateral  lobes  each  divided  into  a 
slender,  finger-like,  membranous  lobe  adjacent  to  apical  ventral  lobe,  a  large,  spreading  membranous  lobe 
and  two  small,  membranous  but  minutely  denticulate  ventral  lobes.  Sclerotized  parts  of  conjunctiva 
consisting  of  these  denticles,  appendages  of  distal  dorsolateral  lobes  and  cup-like  sclerite  protecting  base  of 
vesica.  Vesica  moderately  long,  its  apex  obliquely  truncate.  Ejaculatory  reservoir  apparatus  with  wings  but 
without  straps. 

Head  granulate,  dorsally  with  some  stout  tubercles;  antennae  granulate,  segment  I  also  tuberculate. 
Pronotum  punctate,  granulate  and  tuberculate  with  a  pair  of  large,  somewhat  irregular,  whitish  tubercules 
near  midline  between  posterolateral  angles.  Scutellum  granulate-punctate,  margins  tuberculate.  Thoracic 
pleura  granulate-punctate.  Clavus  and  corium  granulate  and  deeply  punctate,  costal  margin  of  corium  in 
basal  half  tuberculate.  Femora  and  tibiae  granulate  and  tuberculate.  Connexivum  granulate-punctate. 
Abdominal  sterna  granulate,  lateral  margins  tuberculate. 

Pubescence  of  body  and  appendages  of  moderate  length,  semidecumbent,  with  longer,  suberect  or  erect 
hairs  arising  from  larger  tubercles  of  appendages  and  dorsal  surface  of  body,  hairs  arising  from  the 
tubercles  of  head  and  pronotum  particularly  long  and  erect. 

Colour  rufous  brown.  Anterior  midline  of  pronotum,  margins  of  clavi  adjacent  to  scutellum,  distal  veins 
of  corium,  abdominal  spines,  large  areas  of  laterotergites  IV  and  V  and  anterior  and  posterior  margins  of 
VI  and  VII  and  apical  two-thirds  of  posterior  femora  conspicuously  darker.  Bases  of  femora,  first  segment 
of  each  tarsus,  tibiae  except  for  basal  and  apical  annuli,  most  of  clavus  and  basal  half  of  corium  and  spots  on 
darker  parts  of  femora  stramineous.  Hemelytral  membrane  fuscous  hyaline,  some  groups  of  somewhat 
reticulate  veins  dirty  white,  veins  of  anal  angle  conspicuously  white,  the  membrane  surrounding  them  not 
infuscate. 

REMARKS.  The  type-locality  is  cited  only  as  'India  Orientalis';  it  is  assumed  here  that  it  was  in  South  East 
Asia  rather  than  in  the  Indian  subcontinent  on  the  basis  of  the  distribution  of  the  other  material  seen. 


TRIBE  PSEUDOPHLOEINI  163 

DISTRIBUTION.  South  East  Asia.  Hsiao  (1964:  252)  records  this  species  from  Yunnan,  China. 

MATERIAL  EXAMINED 

South  East  Asia  'India  orientalis':  1  $  (holotype)  (Stevens)  (NR). 

Burma:  1  cf ,  Bhamo,  viii.  1885  (Fea,  ex  Distant  coll.)  (BMNH).  Vietnam:  1  $ ,  Dalat,  6  km  S. ,  1400-1500 
m,9.vi.!961  (N.  R.  Spencer)  (BPBM).  Laos:  1  cf,  1  $,  WapikhamthongProv.,KhongSedone,  18.ix.1965 
(native  collector)  and  3. viii.  1965  (Rondon)  (BPBM). 

Hoplolomia  campbellisp.  n. 

(Figs  15-17) 

Length:  cf,  5-7-5-8  mm;  $,6-7  mm. 

Very  similar  to  H.  scabricula  in  size,  form  and  colouring,  including  in  the  structure  of  the  conjunctiva 
(Figs  15, 16)  but  body  slightly  more  depressed  and  slightly  narrower.  Length  of  antennal  segment  I  divided 
by  width  of  head  including  eyes  0 •  85-0 •  93 ;  ratio  of  lengths  of  antennal  segments  about  1  -00 : 0 •  83 : 1  •  22 : 0 •  72 . 
Pronotum  shallowly  declivent,  width  across  apices  of  posterolateral  spines  divided  by  width  of  head 
including  eyes  2-03-2-11.  Scutellum  flat. 

REMARKS.  Very  similar  to  H.  scabricula,  this  species  is  distinguished  from  it  by  the  slightly  more 
dorsoventrally  flattened  body,  this  difference  being  most  apparent  in  the  shallower  slope  of  the  pronotal 
disc  and  the  flat,  not  convex,  scutellum. 

DISTRIBUTION.  Southern  India. 

MATERIAL  EXAMINED 

Holotype  cf ,  India:  Chikaballapura,  iii.1915  (T.  V.  Campbell)  (BMNH). 
Paratypes.  India:  1  cf,  Chikaballapura  (Campbell),  ii.1918;  1  $,  Chikaballapura  ll.i.1918  (BMNH). 

i 

INDOLOMIA  gen.  n. 

Type-species:  Indolomia  conculata  sp.  n. 

Body  robust,  connexivum  widely  expanded. 

Head  slightly  longer  than  pronotum,  with  uniform,  low  granulation.  Antennifers  weakly  divergent, 
outer  apical  processes  triangular,  apically  curved  inwards.  Antennal  segment  I  weakly  clavate,  slightly  but 
distinctly  shorter  than  head;  segment  III  slightly  shorter  than  I,  II  slightly  shorter  than  III,  IV  shortest. 
Bucculae  short,  occupying  only  one-quarter  of  length  of  ventral  midline  of  head.  Rostrum  with  segment  I 
reaching  posteriorly  almost  to  level  of  posterior  margin  of  eye;  segment  II  slightly  shorter  than  I,  III  half  as 
long  as  II,  IV  two-thirds  as  long  as  II. 

Pronotum  strongly  declivent,  its  posterolateral  angles  strongly  produced  anterolaterally,  posterolateral 
margin  straight,  without  prescutellar  spines;  mostly  rather  weakly  and  uniformly  granulate,  with  a  pair  of 
short,  oblique  ridges  near  midline  between  posterolateral  angles,  anterior  and  anterolateral  margins  and 
anterior  half  of  midline  with  some  larger  and  more  prominent  granules.  Scutellum  equilateral,  convex,  its 
apex  slightly  produced  and  swollen.  Meso-  and  metasternum  strongly  sulcate  longitudinally.  Dorsal  ridge 
of  metathoracic  peritreme  shortly  reniform.  Corium  with  costal  margin  weakly  concave  and  apical  margin 
weakly  convex,  apex  not  produced.  Metathoracic  wing  without  antevannal  vein.  Anterior  and  intermedi- 
ate femora  each  with  a  small  subapical  spine  beneath;  posterior  femur  with  two  or  three  major  subapical 
spines  and  an  apical  series  of  tubercles  and  with  a  prominent  basal  tubercle.  Posterior  coxae  separated  by  a 
space  equal  to  about  two-thirds  of  the  width  of  a  coxa. 

Abdominal  sterna  HI- VII  with  posterolateral  angles  triangularly  produced,  lateral  margins  with  very 
obsolete  granulation.  Male  genital  capsule  posteriorly  emarginate,  emargination  filled  by  apices  of 
parameres.  Phallotheca  with  a  broad  ventral  sclerite,  produced  laterally,  and  two  thin,  longitudinal  dorsal 
sclerites.  Phallotheca,  conjunctiva  and  vesica  subequal  in  length.  Conjunctiva  with  sclerotized,  ventrally 
directed  distal  dorsolateral  appendages  and  a  pair  of  sclerites  protecting  coiled  base  of  vesica,  otherwise 
completely  membranous;  dorsomedian  lobe  low,  distal  dorsomedian  lobe  conical,  apical  ventral  lobes 
paired,  weakly  developed,  distal  ventrolateral  lobes  trilobed.  Ejaculatory  reservoir  complex  with  wings 
well  developed,  straps  absent. 

REMARKS.  This  genus  resembles  Hoplolomia  in  that  it  lacks  an  antevannal  vein  in  the  metathoracic  wing 
and  has  a  basal  tubercle  on  the  hind  femur.  Because  of  the  shape  of  the  pronotum,  the  short  pubescence 
and  the  absence  of  prominent  tubercles  from  most  of  the  body  surface  and  appendages,  it  superficially 


164 


W.  R.  DOLLING 


18 


19 


Figs  12-19  12-14,  Hoplolomia  scabricula:  (12)  dorsal  view  of  pronotum;  (13)  ventral  view  of  abdominal 
margin;  (14)  dorsal  view  of  apical  half  of  posterior  femur.  15-17,  H.  campbelli:  (15)  dorsal  view  of 
conjunctiva  and  vesica;  (16)  ventral  view  of  same;  (17)  metathoracic  wing.  18, 19,  Indolomia  conculata: 
(18)  dorsal  view  of  pronotum;  (19)  ventral  view  of  abdominal  margin. 


resembles  some  My  la  species  but  the  differences  in  the  form  of  the  male  genitalia  show  that  the  two  genera 
are  not  closely  releated. 

DISTRIBUTION.  A  single  species,  in  southern  India. 

Indolomia  conculata  sp.  n. 

(Figs  18, 19) 

Length:  d",  7-8  mm;  $  unknown. 

Head  as  long  as  its  width  including  eyes.  Length  of  antennal  segment  I  divided  by  head  width  including 
eyes  0-89.  Ratio  of  lengths  of  antennal  segments  as  l-00:0-89:0-94:0-83.  Specialized  sensory  setae 


TRIBE  PSEUDOPHLOEINI  165 

occupying  about  nine-tenths  of  length  of  segment  IV.  Antennae  without  prominent  granules  or  tubercles. 
Rostral  segment  1 0-66  times  as  long  as  width  of  head  including  eyes;  ratio  of  lengths  of  rostral  segments  as 
l-00:0-89:045:0-64. 

Pr.onotum  (Fig.  18)  with  posterolateral  angles  terminating  in  short  spines;  width  across  apices  of  spines 
2-43  times  width  of  head  including  eyes.  Legs  moderately  long,  posterior  tibia  0-96  times  as  long  as 
posterior  femur.  Left  femur  of  holotype  with  three  major  spines,  the  second  half  as  long  as  the  third  and 
twice  as  long  as  the  first,  right  femur  with  only  two  major  spines,  similar  to  second  and  third  of  left  femur; 
an  apical  series  of  three  spines  present  on  both  femora,  one  tubercle  and  two  granules  present  between  last 
and  penultimate  spines,  a  few  granules  present  on  rest  of  apical  half  of  ventral  surface  of  both  femora. 
Scutellum,  clavus  and  corium  strongly  punctate  and  weakly  granulate  throughout.  Abdominal  sterna  with 
posterolateral  angles  triangularly  produced  but  not  drawn  out  into  spines  (Fig.  19).  Pubescence  of  body, 
antennal  segment  I  and  most  of  femora  short,  semidecumbent;  that  of  rest  of  antennae,  of  tibiae  and  tarsi 
and  parts  of  femora  short  or  rather  short ,  suberect ;  front  of  head  and  some  large  granules  of  pronotum  with 
a  few  longer,  erect  hairs. 

Colour  various  shades  of  brown.  Head  with  midline  and  a  U-shaped  stripe  adjacent  to  eyes  and 
interrupted  by  ocelli  conspicuously  paler.  Pronotum  with  midline  and  posterior  disc  conspicuously  darker, 
the  two  oblique  ridges  of  posterior  disc  conspicuously  paler.  Femora  piceous  with  stramineous  spots. 
Tibiae  stramineous  with  basal  and  apical  annuli  piceous.  Tarsi  piceous,  first  segment  dorsally  stramineous. 
Connexivum  banded  pale  and  dark  brown,  abdominal  dorsum  rufous  brown,  anteriorly  and  anterolateral- 
ly  piceous.  Thoracic  sterna  piceous.  Abdominal  sterna  mottled  pale  brown  and  piceous,  piceous  markings 
tending  to  form  five  longitudinal  stripes.  Hemelytral  membrane  fuscous  hyaline;  veins  darker  brown, 
interrupted  by  milky  spots;  anal  angle  of  membrane,  including  veins,  milky. 

REMARKS.  Only  the  holotype,  which  is  complete  but  slightly  crushed,  is  known. 

MATERIAL  EXAMINED 
Holotype  cf ,  India:  Anaimalai  Hills,  Cinchuna,  3500  ft  [1067  m],  v.1976  (T.  Nathan)  (BMNH). 

PARAM YLA  Linnavuori  stat.  n. 

Myla  (Paramyla)  Linnavuori,  1971:  177-178.  Type-species:  Myla  suspecta  Schouteden,  by  original 
designation. 

Body  depressed,  narrowly  ovate,  not  conspicuously  tuberculate  or  hispid,  connexivum  moderately 
expanded. 

Head  slightly  shorter  than  pronotum,  slightly  longer  than  antennal  segment  I,  dorsally  granulate  with 
two  conspicuous,  smooth,  non-granulate  lines  arising  between  ocelli  and  diverging  anteriorly,  reaching 
half-way  to  antennifers,  and  a  shorter,  smooth  line  along  outer  margin  of  each  ocellus.  Antennifers 
divergent,  outer  apical  processes  of  antennifers  short,  porrect.  Antennal  segment  I  longest,  weakly  but 
rather  abruptly  clavate,  II  shortest,  III  next  shortest;  I-III  densely  granulate,  granules  of  I  larger;  IV  with 
specialized  sensory  area  occupying  between  two-thirds  and  four-fifths  of  its  length.  Bucculae  occupying 
about  one-quarter  of  ventral  midline  of  head.  Rostrum  with  segment  I  distinctly  longest,  II  longer  than  IV, 
III  shortest,  apex  of  rostrum  at  rest  reaching  to  disc  of  mesosternum. 

Pronotum  shallowly  declivent,  posterolateral  angles  scarcely  prominent,  bearing  a  short,  triangular 
tooth,  lateral  margins  granulate,  disc  granulate  to  tuberculate,  posterior  margin  weakly  convex,  prescutel- 
lar  spines  well  developed.  Scutellum  flat,  equilateral,  apex  slightly  or  strongly  elevated.  Mesosternum 
sulcate,  metasternum  convex  with  slight  longitudinal  groove  in  anterior  one- third.  Dorsal  ridge  of 
metathoracic  peritreme  reniform.  Metathoracic  wing  without  antevannal  vein.  All  femora  granulate, 
anterior  two  pairs  with  or  without  a  subapical  spine  beneath,  posterior  femur  with  a  moderately 
well-developed  basal  tubercle  and  with  two  or  three  major  subapical  spines  with  only  granules  between 
them  and  an  apical  series  of  two  to  four  tubercles.  Posterior  tibiae  more  than  0-90  times  length  of  posterior 
femora. 

Abdominal  sternites  with  their  posterolateral  angles  not  produced,  their  densely  granulate  lateral 
margins  forming  a  single,  smooth  curve.  Genital  capsule  (Fig.  21)  short,  posteriorly  emarginate,  emargina- 
tion  filled  by  apices  of  parameres  (Fig.  22). 

REMARKS.  Members  of  this  genus  are  readily  recognized  among  tropical  Pseudophloeinae  by  their  strongly 
depressed  and  largely  non-tuberculate  body.  They  have  more  the  appearance  of  some  Palaearctic  genera, 
but  differ  from  these  in  the  absence  of  an  antevannal  vein  in  the  metathoracic  wing.  Linnavuori  (1971) 
included  Myla  niokoensis  Schouteden  in  his  new  subgenus  Paramyla  but  the  male  genitalia  of  niokoensis 
are  typical  of  the  genus  Myla  and  the  resemblances  between  niokoensis  and  suspecta  are  confined  to  the 


W.  R.  DOLLING 


Figs  20-26  Paramyla  species.  20,  australis,  dorsal  view  of  body  with  hemelytra  at  rest  and  posterior  femur 
and  tibia.  21-26,  suspecta:  (21)  posterior  view  of  male  genital  capsule  with  parameres;  (22)  dorsomedial 
view  of  left  paramere;  (23)  lateral  view  of  phallotheca,  conjunctiva  and  vesica;  (24)  ventral  view  of  same; 
(25)  dorsal  view  of  head  and  pronotum;  (26)  dorsal  view  of  posterior  femur. 


superficial  characters  of  a  lack  of  abdominal  spines,  very  short  pronotal  spines  and  very  short  pubescence. 
DISTRIBUTION.  Mountainous  regions  of  central  Africa  and  South  Africa. 

Key  to  species 

1    Pronotum  with  prescutellar  spines  broad,  triangular;  hairs  of  pubescence  longer  than  distance 

between  adjacent  hair  insertions.  (Mountains  of  central  Africa) suspecta  (p.  166) 

Pronotum  with  prescutellar  spines  slender;  hairs  of  pubescence  shorter  than  distance  between 
adjacent  insertions.  (South  Africa) australis(p.  167) 

Paramyla  suspecta  (Schouteden)  comb.  n. 
(Figs  21-26) 

Myla  suspecta  Schouteden,  1938:  294.  Holotype  cf ,  ZAIRE  (MRAC)  [examined]. 
Myla  (Paramyla)  suspecta  (Schouteden);  Linnavuori,  1971:  177-178. 

Length:  cf,  7-4-7-6  mm;  $,7-7-8-0 mm. 

Antennal  segment  I  with  two  different  sizes  of  granulation,  its  length  approximately  equal  (0-89-1-01 
times)  to  width  of  head  including  eyes;  ratio  of  lengths  of  antennal  segments  about  l-00:0-67:0-81:0-85; 


TRIBE  PSEUDOPHLOEINI  167 

specialized  sensory  area  of  segment  IV  occupying  about  four-fifths  of  its  length.  Ratio  of  lengths  of  rostral 
segments  about  1-00:0-80:0-45:0-70. 

Pronotum  (Fig.  25)  with  disc  granulate;  lateral  margins  shallowly  concave;  posterolateral  angles  scarcely 
elevated  above  general  level  of  posterior  disc,  width  across  apices  of  blunt  posterolateral  spines  1-73-1-82 
times  width  of  head  including  eyes;  prescutellar  angles  with  triangular  spines.  Scutellum  with  apex  slightly 
elevated.  Anterior  and  intermediate  femora  without  subapical  spines  or  tubercles,  posterior  femur  (Fig. 
26)  with  two  major  spines,  the  proximal  one  about  half  as  long  as  the  other,  with  only  granules  between 
them,  and  an  apical  series  of  two  or  three  tubercles.  Corium  with  costal  margin  markedly  convex. 

Male  with  phallotheca  short,  consisting  of  a  single  ventral  sclerite  extending  laterally  almost  to  meet  two 
rather  less  well-sclerotized  dorsal  sclerites.  Conjunctiva  (Figs  23,  24)  with  dorsomedian  lobe  large,  its 
lateral  angles  prominent,  distal  dorsomedian  lobe  low,  scarcely  developed,  distal  dorsolateral  lobes 
finger-like,  wholly  membranous  and  without  sclerotized  appendages,  apical  ventral  lobes  absent,  distal 
ventrolateral  lobes  each  divided  into  three:  a  dorsal,  wholly  membranous  part,  a  ventral,  largely 
sclerotized  part,  denticulate  along  its  anterior  edge,  and  a  narrow,  denticulate  lobe  situated  anterior  and 
medial  to  the  two  larger  parts.  Vesica  stout,  rather  long,  protected  at  base  by  a  pair  of  sclerites.  Ejaculatory 
reservoir  complex  with  wings  long,  L-shaped,  straps  absent. 

Head  granulate  throughout,  granules  larger  along  midline  of  tylus,  gular  region  punctate.  Antennal 
segment  I  with  small,  densely  packed  granules  and  some  larger,  outstanding  granules,  segments  II-III  with 
very  small  granules.  Pronotal  disc  punctate,  with  scattered  granules,  anterior  half  with  some  larger 
granules  or  low  tubercles,  lateral  margins  with  larger  granules;  scutellum  weakly  granulate-punctate; 
thoracic  sterna  and  pleura  punctate;  femora  and  tibiae  minutely  granulate.  Clavus  and  corium  punctate 
throughout,  punctures  of  clavus  larger,  of  apex  of  corium  smaller  than  average  size  of  those  on  disc  of 
corium,  veins  of  clavus  and  corium  granulate.  Abdominal  sternites  and  laterotergites  minutely  granulate, 
laterotergites  also  minutely  punctate. 

Insect  clothed  in  short,  semidecumbent  pubescence,  the  hairs  longer  than  the  distances  between  their 
insertions;  tubercles  and  larger  granules  of  head  and  pronotum  with  longer,  erect  hairs. 

Colour  largely  mid-brown.  Membrane  of  hemelytra  infuscate  between  veins,  veins  brown,  with  short, 
paler  interruptions.  Antennal  segment  IV,  femora,  especially  posterior  pair,  and  mesosternum  dark  brown 
to  black;  tibiae,  except  for  basal  and  apical  annuli,  paler  brown. 

REMARKS.  This  species  seems  to  be  one  of  the  tropical  Pseudophloeinae  most  highly  adapted  for  life  on  the 
ground.  The  unusual  form  of  the  conjunctiva,  lacking  apical  ventral  lobes  and  sclerotized  appendages  to 
the  distal  dorsolateral  lobes  and  with  very  strongly  developed  distal  ventrolateral  lobes,  is  presumably 
characteristic  for  the  genus,  and  sets  it  somewhat  apart  from  other  genera. 

DISTRIBUTION.  Highlands  of  central  Africa. 

MATERIAL  EXAMINED 

Zaire:  1  cf  (holotype),  Ituri,  Nioka,  vii.1934  (/.  V.  Leroy)  (MRAC). 

Zaire:  4  cf,  4  $,  data  as  holotype  except  that  one  male  and  two  females  have  fuller  date:  7. vii.1934 
(probably  all  paratypes);  1  cf,  Ituri,  de  Buba  a  Petro  (Nizi),  23.iii.1929  (A.  Collarf)  (paratype);  1  $, 
Kibali-Ituri,  Nioka,  xii.1952  (J.  Hecq)  (MRAC).  Malawi:  1  cf ,  Masuku  Mts,  6000-7000  ft  [1800-2100  m], 
vii.l896(A  Whyte)  (BMNH). 

Paramyla  australis  sp.  n. 

(Fig.  20) 

Length:  cf ,  unknown;  $,7-5  mm. 

Antennae  with  segment  I  proportionately  longer  and  segment  IV  much  shorter  than  in  P.  suspecta,  ratio 
of  lengths  of  segments  as  l-00:0-71:0-87:0-65;  granulation  of  segment  I  uniform,  segment  IV  with  junction 
of  specialized  sensory  area  and  area  of  normal  pubescence  oblique,  specialized  area  occupying  between 
three-quarters  and  two-thirds  of  its  length.  Body  form  (Fig.  20)  slightly  narrower  than  that  of  suspecta; 
width  of  pronotum  (Fig.  20)  across  apices  of  the  blunt,  triangular  posterolateral  spines  1-72  times  width  of 
head  including  eyes;  prescutellar  spines  slender;  posterolateral  angles  rather  strongly  elevated  above  level 
of  posterior  disc;  lateral  margins  strongly  concave.  Scutellum  with  apex  strongly  raised.  Corium  with  costal 
margin  only  weakly  convex.  Intermediate  femur  with  a  small  subapical  spine  beneath;  posterior  femur 
(Fig.  20)  with  three  major  spines,  of  which  the  first  is  very  small  and  the  second  about  three-quarters  the 
length  of  the  last,  a  single  tubercle  between  these  two  biggest  spines  and  an  apical  series  of  four  tubercles. 

Sculpture  of  body,  especially  pronotum,  rather  more  pronounced  than  in  suspecta;  pronotal  disc 
tuberculate.  Pubescence  of  body  and  appendages  very  short,  semidecumbent,  white,  the  hairs  distinctly 


168  W.  R.  DOLLING 

shorter  than  the  distances  between  their  insertions;  tarsi  and  apices  of  tibiae  with  longer,  suberect 
pubescence  but  body  without  longer,  suberect  hairs. 

REMARKS.  This  species  diverges  less  from  the  typical  body  form  of  the  subfamily  than  does  its  congener;  the 
body  is  less  flattened  and  there  are  more  femoral  spines.  In  contrast,  the  pubescence  is  abnormally  short  in 
the  new  species. 

DISTRIBUTION.  Known  only  from  the  type-locality  in  South  Africa. 

MATERIAL  EXAMINED 

Holotype  £,  South  Africa:  Cape  Province,  Cape  Peninsula,  Red  Hill,  30.U968  (/.  A.  &  S.  Slater,  T. 
Schuh,  M.  H.  Sweet)  (J.  A.  Slater  collection). 

PSILOLOMIA  Breddin 

Psilolomia  Breddin,  1909:  292.  Type-species:  Psilolomia  brevitibialis  Breddin,  by  monotypy. 
Neohoplolomia  Distant,  1913:  284.  Type-species:  Neohoplolomia  typica  Distant,  by  monotypy.  Syn.  n. 
Psilocoris  Hsiao,  1964:  251,  259.  Type-species:  Psilocoris  clavipes  Hsiao,  by  original  designation.  Syn.  n. 

Body  oblong,  not  depressed,  connexivum  rather  narrowly  to  moderately  expanded,  body  not  conspicu- 
ously spinose,  tuberculate  or  hairy. 

Head  about  as  long  as  pronotum,  dorsally  granulate  to  rather  weakly  tuberculate.  Antennifers 
divergent,  their  outer  apical  processes  short,  broad,  porrect.  Antennae  with  segment  I  very  weakly  clavate, 
granulate,  without  conspicuously  outstanding  granules  or  tubercules;  segments  II  and  III  never  of 
conspicuously  unequal  length,  minutely  granulate.  Bucculae  occupying  about  one-quarter  of  ventral 
midline  of  head .  Rostrum  with  segment  I  much  the  longest ,  II  to  IV  subequal  in  length ,  apex  of  IV  reaching 
to  meso-metasternal  suture. 

Pronotum  moderately  declivent,  its  posterolateral  angles  not  or  rather  weakly  elevated  and  not  or  rather 
weakly  produced  laterally,  posterolateral  spines  short,  blunt  or  acute  and  arising  abruptly  from  the  angles, 
directed  laterally;  posterior  margin  straight,  without  humeral  spines;  disc  and  lateral  margins  with  large 
granules  or  tubercles.  Scutellum  flat  or  weakly  convex,  equilateral,  anterior  angles  prominent,  apex 
elevated  as  a  minute,  whitish  blob.  Mesosternum  shallowly  sulcate;  metasternum  almost  flat  to  distinctly 
convex,  anteriorly  concavely  emarginate.  Metathoracic  peritreme  (Fig.  38)  with  dorsal  ridge  bilobed,  the 
anterior  lobe  much  the  larger.  Clavus  and  corium  punctate  throughout,  corium  less  heavily  so  apically  and 
with  apex  not  produced.  Antevannal  vein  present  in  metathoracic  wing  (Fig.  39)  but  sometimes  weak. 
Anterior  and  intermediate  femora  with  zero  to  two  small  subapical  spines  beneath,  posterior  femur  with 
three  major  subapical  spines,  some  smaller  ones  and  an  apical  series;  tubercle  at  base  of  posterior  femur 
small,  sometimes  obsolete.  Posterior  coxae  not  approximated. 

Abdominal  sterna  III- VII  with  posterolateral  angles  not  or  slightly  produced  (maximum  development  in 
P.  brunneofusca,  Fig.  42).  Spermatheca  with  bulb  lunate,  duct  short,  not  convoluted.  Male  genital  capsule 
posteriorly  emarginate,  emargination  filled  by  apices  of  parameres.  Conjunctiva  with  dorsomedian  lobe 
large,  distal  dorsomedian  lobe  smaller,  both  membranous;  distal  dorsolateral  lobes  slender,  their  posterior 
faces  supported  for  much  or,  usually,  all  of  their  length  by  sclerites  continuous  with  or  articulating  with 
wings  of  ejaculatory  reservoir,  which  lacks  straps;  no  sclerites  protecting  base  of  vesica;  apical  ventral  lobe 
obsolete  or  transverse,  sometimes  weakly  bilobed  but  never  divided  into  two  globular  lobes;  distal 
ventrolateral  lobes  membranous;  ventral  wall  of  conjunctiva  with  a  pair  of  posteriorly  diverging,  toothed, 
sclerotized  strips,  these  weakly  developed  or  absent  in  the  smallest  three  species. 

Pronotum,  scutellum  and  thoracic  pleura  granulate-punctate,  veins  of  hemelytra  with  occasional  small 
granules,  femora,  tibiae,  abdominal  sternites  and  laterotergites  granulate.  Pubescence  colourless  or 
amber,  simple,  mostly  rather  short,  semidecumbent  to  suberect,  some  species  with  long,  erect  hairs  on 
pronotum  and  dorsum  of  head;  pubescence  never  scale-like  or  tomentose. 

REMARKS.  This,  one  of  the  largest  genera  of  the  tribe,  lacks  any  striking  distinguishing  features.  The 
type-species  of  Psilolomia  and  Neohoplolomia  are  synonymized  here.  P.  brevitibialis,  the  type-species  of 
Psilolomia,  is  unique  in  that  its  vesica  is  about  twice  as  wide  as  those  of  the  other  species  examined  and  the 
distal  dorsolateral  lobes  of  the  conjunctiva  are  sclerotized  for  only  about  half  their  length.  It  shares  with 
only  P.  parva  both  the  absence  of  ventral,  toothed,  sclerotized  strips  from  the  conjunctiva  and  the 
possession  of  an  antennal  segment  II  that  is  shorter  than  segment  III;  these  are  the  only  two  species  in 
which  some  of  the  dorsal  granules  of  the  head  are  developed  into  tubercles  that  are  higher  than  wide.  The 
conjunctiva  of  P.  amphrysia,  the  third  small  species,  has  very  weakly  developed  ventral  sclerotized  strips 
and  thus  forms  a  link  between  the  other  two  and  the  large  species  of  the  genus.  These  differences  do  not 
seem  of  sufficient  value  to  warrant  the  separation  of  P.  brevitibialis  or  of  P.  brevitibialis  plus  P.  parva  from 


TRIBE  PSEUDOPHLOEINI 


169 


Figs  27-35    Psilolomia  species,  outline  dorsal  view  of  right  half  of  pronotum  of  27,  amphrysia;  28, 
nigeriensis;  29,  lamottei;  30,  brevitibialis;  31,parva;  32,  vulgaris;  33,  brunneofusca;  34,  steeleae;  35,  lata. 


the  remaining  species  at  the  generic  level.  Linnavuori  (1970)  placed  the  two  African  species  known  to  him 
in  Mevaniomorpha,  which  is  undoubtedly  close  to  Psilolomia  and  might  be  regarded  as  a  derivative  of  it 
with  more  pronounced  pronotal  and  abdominal  spines. 

DISTRIBUTION.  Afrotropical  and  Oriental  regions;  wholly  continental,  with  no  insular  records. 

Key  to  species 

1  Side  of  head  above  each  antennifer  with  a  large  tubercle  almost  as  large  as  an  eye  (Figs  36, 37) . 

(India) d/spar(p.  174) 

-  Head  without  such  tubercles 2 

2  Posterolateral  angles  of  pronotum  bearing  acute  spines  that  are  longer  than  their  basal  width 

(Figs  1,27, 30, 31, 33) .....          3 

-  Posterolateral  angles  of  pronotum  with  usually  blunt  spines  no  longer  than  their  basal  width 

(Figs  28, 29, 32, 34, 35) 7 


170  W.  R.  DOLLING 

3  Larger  insects:  length  of  male  7-6  mm  or  more;  length  of  female  7-9  mm  or  more.  Antennal 

segment  I  as  long  as  or  longer  than  width  of  head  including  eyes 4 

Smaller  insects:  length  of  male  7-2  mm  or  less;  length  of  female  7-6  mm  or  less.  Antennal 
segment  I  not  as  long  as  width  of  head  including  eyes 5 

4  Coloration  ochreous  yellow.  (India,  Sri  Lanka) pundaloyae (p.  177) 

Coloration  ferruginous  or  fuscous  brown.  (S.  China  to  Malaya) 

brunneofusca(p.  Ill),  clavipes(p.  178) 

5  Antennal  segment  II  longer  than  segment  III.  (Africa) amphrysia  (p.  170) 

Antennal  segment  II  shorter  than  segment  III.  (Asia) 6 

6  Pronotum  (Fig.  31)  with  lateral  margins  distinctly  concave;  abdominal  sternites  V  and  VI  with 

posterolateral  angles  produced  into  short,  acute  spines.  (Laos,  Thailand) parva  (p.  174) 

Pronotum  (Fig.  30)  with  lateral  margins  almost  straight;  abdominal  sternites  V  and  VI  with 
posterolateral  angles  right-angled.  (India,  Sri  Lanka) brevitibialis  (p.  172) 

7  Pronotum  with  lateral  margins  (Figs  32,  34,  35)  bearing  coarser  tubercles  and  granules  and 

longer  and  straighter  pubescence.  (Asia) 

Pronotum  with  lateral  margins  (Figs  28,  29)  bearing  finer  tubercles  and  shorter,  more  curved 
pubescence.  (Africa) 10 

8  Pronotum  with  lateral  margins  (Fig.  35)  strongly  concave  just  anterior  to  posterolateral  angles. 

(Thailand) Ma  (p.  179) 

Pronotum  with  lateral  margins  (Figs  32, 34)  weakly  concave 9 

9  Antennal  segment  I  longer  than  width  of  head  including  eyes.  General  coloration  ferruginous 

brown.  (Burma) steeleae(p.  178) 

Antennal  segment  I  shorter  than  width  of  head  including  eyes.  General  coloration  ochreous 
yellow.  (India,  Sri  Lanka) vulgaris(p.  175) 

10  Antennal  segment  I  not  longer  than  width  of  head  including  eyes.  Pronotum  with  lateral 

margins  (Fig.  28)  almost  straight.  (Nigeria,  Central  African  Republic) nigeriensis  (p.  171) 

Antennal  segment  I  longer  than  width  of  head  including  eyes.  Pronotum  with  lateral  margins 
(Fig.  29)  distinctly  concave 11 

11  Larger  insects:  length  of  male  9-9  mm.  (Sudan) ferruginea(p.  172) 

Smaller  insects:  length  of  male  9-0  mm;  length  of  female  9-7  mm.  (Guinea,  Nigeria)     Iamottei(p.  171) 

Psilolomia  amphrysia  (Linnavuori)  comb.  n. 
(Figs  27,  49) 

Mevaniomorpha  amphrysius  Linnavuori,  1970:  43-45.  Holotype  C?,  IVORY  COAST:  Lamto,  Toumodi, 
1-5. vi.  1962  (Gillori)  (R.  Linnavuori  coll.)  [not  examined]. 

Length:  d",  6-6-7-2 mm;  $,7-2-7-6 mm. 

Head  dorsally  granulate.  Antennifers  distinctly  but  not  strongly  divergent.  Length  of  antennal  segment  I 
divided  by  width  of  head  including  eyes  0-88-1-00.  Ratio  of  lengths  of  antennal  segments  in  male  about 
l-00:0-78:0-73:0-82,  in  female  about  l-00:0-77:0-71:0-74.  Ratio  of  lengths  of  rostral  segments  in  male 
about  l-00:0-65:0-50:0-73,  in  female  about  1-00:0-58:0-52:0-61. 

Pronotum  (Fig.  27)  with  lateral  margins  slightly  concave,  bearing  small  tubercles  that  are  slightly  longer 
than  wide;  posterolateral  spines  acute,  longer  than  their  basal  width;  width  across  apices  of  spines  divided 
by  width  of  head  including  eyes  1-80-2-07.  Metasternum  moderately  convex.  Anterior  and  intermediate 
femora  with  or  without  one  small,  subapical  spine  beneath,  posterior  femur  with  three  major  spines,  three 
or  four  minor  spines  between  the  second  and  third  and  an  apical  series  of  four.  Length  of  posterior  tibia 
about  0-7  times  length  of  posterior  femur. 

Abdominal  sternites  III-VII  with  posterolateral  angles  acute,  slightly  prominent.  Conjunctiva  with 
distal  dorsolateral  lobes  (Fig.  49)  short,  sclerotized  throughout  their  length;  ventral  sclerotized  strips 
weakly  developed. 

Pubescence  of  most  of  body  and  of  hemelytra  short,  curved,  semidecumbent;  that  of  antennae  and  legs 
straighter,  suberect;  head  also  with  some  longer,  suberect  hairs  arising  from  tubercles  and  larger  granules; 
declivent  part  of  pronotum  also  with  pubescence  of  erect  hairs  about  as  long  as  diameter  of  an  eye. 

Colour  sordid  yellow,  apical  half  of  posterior  femur  heavily  marked  piceous;  scattered,  small  piceous 
markings  of  greater  or  less  extent  present  elsewhere  on  body  and  appendages,  especially  abdominal 
laterotergites  and  sternites.  Granules  of  appendages  usually  piceous. 

REMARKS.  This  is  the  smallest  of  the  African  species  and  the  only  one  of  them  with  long  pronotal 
pubescence  (about  as  long  as  diameter  of  an  eye);  its  pronotal  posterolateral  spines  are  longer  than  those  of 


TRIBE  PSEUDOPHLOEINI  171 

other  African  species.  It  differs  from  the  two  small  Asian  species  in  having  the  second  antennal  segment 
longer  than  the  third,  not  shorter. 

DISTRIBUTION.  West  Africa  and  Zambia. 

MATERIAL  EXAMINED 

Ivory  Coast:  1  cf ,  Lamto,  3.iii.l967  (Gillon),  1  $,  Lamto,  7-10.ii.1962  (Gillon)  (R.  Linnavuori  coll; 
paratypes);  6  cf ,  Lamto,  various  dates  26.vi.1962  to  21.vii.1964  (Gillon)  (BMNH).  Nigeria:  1  cf ,  Samaru, 
Zaria,  in  dry  plant  detritus,  30.U955  (M.G.E.);  1  cf,  1  $,  Ilorin,  under  dead  leaf ,  19.iii.1955  (A/.G.E.);3 
$ ,  Gombe,  Matzoro  Lakes,  i.1929  (Lloyd)  (BMNH).  Zambia:  1  cf ,  Chisinga  Plateau,  Kalungwisi  District, 
4500  ft  (1350  m),  25.ix.1908  (Neave)  (UM). 

Psilolomia  nigeriensis  sp.  n. 

(Figs  28,  57) 

Length:  cf ,  7-9-9-3  mm;  $,8-7-9-1  mm. 

Head  dorsally  granulate.  Antennifers  distinctly  but  not  very  abruptly  divergent.  Length  of  antennal 
segment  I  divided  by  width  of  head  including  eyes  0-86-1-00.  Ratio  of  lengths  of  antennal  segments  in 
northern  males  about  l-00:0-89:0-78:0-93;  in  southern  males  about  l-00:0-93:0-80:0-82;  in  northern 
females  about  l-00:0-93:0-82:0-86;  in  southern  females  about  l-00:0-93:0-82:0-82.  Ratio  of  lengths  of 
rostral  segments  about  1.00:0-57:0-58:0-55. 

Pronotum  (Fig.  28)  with  lateral  margins  very  shallowly  concave,  almost  straight,  bearing  small  granules; 
posterolateral  angles  each  with  a  short,  triangular  spine;  width  across  apices  of  spines  divided  by  width  of 
head  including  eyes  1-94:2-02.  Metasternum  rather  strongly  convex.  Anterior  and  intermediate  femora 
each  without  or  with  one  small,  subapical  spine  beneath;  posterior  femur  with  three  major  spines,  rarely 
with  a  small  spine  preceding  them,  three  or  four  minor  spines  between  the  penultimate  and  last  major 
spines  and  an  apical  series  of  four  or  five.  Length  of  posterior  tibia  divided  by  length  of  posterior  femur 
0-82-0-92. 

Abdominal  sternites  III-VII  with  posterolateral  angles  acute  and  slightly  prominent.  Conjunctiva  with 
ventral  toothed  sclerotized  strips  well-developed,  distal  dorsolateral  lobes  (Fig.  57)  sclerotized  throughout 
their  length  but  less  strongly  so  apically  and  ventrally,  dorsomedian  lobe  bearing  a  narrow,  membranous, 
apically  bifid,  median  process;  vesica  longer  than  in  any  other  species  of  the  genus. 

Pubescence  of  most  of  body  and  of  hemelytra  short,  curved,  semidecumbent;  of  antennae  and  legs 
suberect,  longer  and  more  erect  ventrally  on  femora;  head  with  uniform,  short,  semidecumbent  pubesc- 
ence; declivent  part  of  pronotum  with  longer,  less  curved,  suberect  pubescence  distinctly  shorter  than 
diameter  of  eye. 

Colour  sordid  yellow-  ochreous,  usually  with  a  slight  ferruginous  tinge  and  with  piceous  markings  along 
apical  margin  of  corium,  thoracic  and  abdominal  sterna  and  laterotergites  and  apical  half  of  posterior 
femur;  all  tibiae  with  basal  and  apical  annuli  fuscous. 

REMARKS.  This  species  differs  from  the  other  African  species  in  its  almost  straight  lateral  pronotal  margins. 
It  is  very  similar  to  the  Asian  P.  vulgaris  but  lacks  suberect  pubescence  along  the  midline  of  the  head  and 
differs  in  the  shape  of  the  sclerotized  distal  dorsolateral  lobes  of  the  conjunctiva. 

DISTRIBUTION.  West  Africa. 

MATERIAL  EXAMINED 

Holotype  cf ,  Nigeria:  U.C.  Ibadan,  on  leaf  of  Vigna  unguiculata,  31.1.1955  (G.  H.  Caswelf)  (BMNH). 

Paratypes.  Nigeria:  1  cf ,  data  as  holotype  (Ibadan  University) ;  1  $,  Enugu,  30.x. 1955  (Bechyne,  Exped. 
Mus.  G.  Frey);  1  cf ,  1  $ ,  Zaria,  Samaru,  at  light,  12.vii.1966  (J.  Deeming);  1  cf ,  2  $ ,  Samaru,  Zaria,  in  dry 
plant  detritus,  dry  grass  and  plant  bases,  30.i.l955  (M.G.E.)  (BMNH).  Central  African  Republic:  1  cf , 
Bambari,  u.v.,  1964  (G.  Pierrard)  (MRAC). 

Psilolomia  lamottei  ( Villiers)  comb.  n. 

(Figs  29,  54) 
Acanthomia  lamottei  Villiers,  19506:  654-655.  Holotype  $,  GUINEA  (MNHN)  [examined]. 

Length:  cf,9-0mm;  $,9-7 mm. 

Head  dorsally  granulate.  Antennifers  strongly  divergent.  Length  of  antennal  segment  I  divided  by  width 
of  head  including  eyes  about  1-1.  Ratio  of  lengths  of  antennal  segments  about  l-00:0-84:0-78:0-76.  Ratio  of 
lengths  of  rostral  segments  about  1 -00:0-58:0-61 :0-58. 


172  W.  R.  DOLLING 

Pronotum  (Fig.  29)  with  anterolateral  margins  concave,  granulate;  posterolateral  angles  prominent, 
bearing  broadly  triangular  posterolateral  spines;  width  across  apices  of  spines  divided  by  width  of  head 
including  eyes  2-05  in  male,  1-93  in  female.  Metasternum  rather  strongly  convex.  Anterior  femur  with  one, 
minute,  subapical  spine  beneath,  intermediate  femur  with  one  small  spine,  posterior  femur  with  three 
major  spines  (preceded  by  a  small  spine  in  holotype)  with  three  minor  spines  between  the  penultimate  and 
last  major  spines  and  an  apical  series  of  four;  tubercle  at  base  of  posterior  femur  obsolete.  Length  of 
posterior  tibia  divided  by  length  of  posterior  femur  0-84-0-85. 

Abdominal  sternites  V  and  VI  with  posterolateral  angles  slightly  prominent,  slightly  more  acute  than 
right-angled;  those  of  VII  angled  at  about  85°,  slightly  prominent.  Vesica  long  and  slender;  conjunctiva 
with  ventral  toothed  strips  well  developed;  distal  dorsolateral  lobes  (Fig.  54)  sclerotized  throughout,  not 
apically  angled  or  expanded. 

Pubescence  rather  short,  suberect  to  semidecumbent,  of  uniform  length,  no  erect  hairs  present  even  on 
dorsal  midline  of  head. 

Coloration  of  holotype  from  Guinea  sordid  yellow  with  slight  rufous  tinge,  antennal  segment  IV, 
tubercles  of  all  appendages,  basal  annulus  of  posterior  tibia  and,  less  conspicuously,  of  other  tibiae,  meso- 
and  metasterna  and  extensive  mottling  on  abdominal  sterna,  laterotergites  and  apical  half  of  posterior 
femur  piceous  or  black.  Male  from  Nigeria  generally  ferruginous  without  black  or  piceous  markings  except 
for  those  of  abdomen  and  thoracic  sterna. 

REMARKS.  There  is  a  close  resemblance  in  structure,  though  not  in  colour,  between  the  two  specimens 
examined.  This  species  resembles  P.  ferruginea  in  having  the  first  antennal  segment  longer  than  the  width 
of  the  head  across  the  eyes  and  in  the  prominent  posterolateral  angles  of  the  pronotum,  but  it  is  smaller  and 
has  fewer  minor  spines  on  the  posterior  femur.  The  two  species  are  evidently  closely  related. 

DISTRIBUTION.  Uplands  of  West  Africa. 

MATERIAL  EXAMINED 

Guinea:  1  $  (holotype),  Nimba,  ii.vi.42  (M.  Lamotte)  (MNHN). 
Nigeria:  1  tf,  Jos,  14.X.1955  (Bechyne,  Exped.  Mus.  G.  Frey)  (BMNH). 

Psilolomia  ferruginea  (Linnavuori)  comb.  n. 

Mevaniomorpha  ferruginea  Linnavuori,  1970:  43  [in  key];  1978:  36-37  [full  description].  Holotype  cf, 
SUDAN  (R.  Linnavuori  coll.)  [examined]. 

Length:  cf ,  9-9  mm;  $,  unknown. 

Head  dorsally  granulate.  Antennifers  strongly  divergent.  Length  of  antennal  segment  I  divided  by  width 
of  head  including  eyes  about  1-2.  Ratio  of  lengths  of  antennal  segments  as  l-00:0-90:0-77:0-63.  Ratio  of 
lengths  of  rostral  segments  as  1 -00:0-71 :0-57:0-68. 

Lateral  margins  of  pronotum  very  distinctly  concave,  bearing  granules  about  as  high  as  wide;  postero- 
lateral angles  prominent,  bearing  short,  broad  posterolateral  spines.  Metasternum  rather  strongly  convex. 
Anterior  and  intermediate  femora  each  with  a  strong  subapical  spine  beneath;  posterior  femur  with  three 
major  spines,  four  minor  spines  between  the  first  pair,  three  or  four  between  the  middle  and  apical  major 
spines  and  an  apical  series  of  five. 

Abdominal  sternites  HI- VII  with  posterolateral  angles  more  acute  than  right-angled,  slightly  promin- 
ent. 

Coloration  generally  ferruginous. 

REMARKS.  This  species  is  closely  similar  to  P.  lamottei.  It  is  the  largest  species  of  the  genus. 
DISTRIBUTION.  Known  only  from  the  type-locality  in  the  foothills  of  the  Ethiopian  massif. 

MATERIAL  EXAMINED 
Sudan:  1  cf  (holotype),  Blue  Nile,  Ingessana  Mts,  17-22. xi. 1962  (R.  Linnavuori)  (R.  Linnavuori  coll.). 

Psilolomia  brevitibialis  Breddin 
(Figs  30, 44, 50) 

Psilolomia  brevitibialis  Breddin,  1909:  293.  LECTOTYPE  $,  SRI  LANKA  (IP),  here  designated  [ex- 
amined] . 

Neohoplolomia  typica  Distant,  1913:  284-285.  LECTOTYPE  cf,  INDIA  (BMNH),  here  designated 
[examined].  Syn.  n. 


TRIBE  PSEUDOPHLOEINI 

Length:  cf,  5-8-6-4  mm;  $,6-4-7-3  mm. 

Head  dorsally  granulate,  tuberculate  throughout  vertex  and  midline.  Antennifers  distinctly  divergent. 
Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes  0-78-0-81.  Ratio  of  lengths  of 
antennal  segments  about  l-00:0-79:0-96:0-83.  Ratio  of  lengths  of  rostral  segments  about  1-00:0-50:0-54: 
0-54.  Head  and  pronotum  together  constituting  less  than  0-39  (male)  or  0-35  (female)  of  total  length. 

Pronotum  (Fig.  30)  with  lateral  margins  straight,  tuberculate;  posterolateral  angles  not  prominent; 
posterolateral  spines  short,  acute;  width  across  apices  of  spines  divided  by  width  of  head  including  eyes 
1-82-1-92.  Metasternum  moderately  convex.  Anterior  and  intermediate  femora  each  without  or  with  one 
small  subapical  spine  beneath;  posterior  femur  with  three  major  spines,  the  first  about  half  as  long  as  the 
others,  one  to  three  minor  spines  between  the  two  large  ones  and  an  apical  series  of  four.  Length  of 
posterior  tibia  divided  by  length  of  posterior  femur  about  0-74. 

Abdominal  sternites  III-VII  with  posterolateral  angles  right-angled,  not  prominent.  Male  conjunctiva 
(Fig.  44)  with  apical  ventral  lobes  obsolete  and  ventral  sclerotized  strips  absent;  distal  dorsolateral  lobes 
(Fig.  50)  short,  supported  by  sclerotizations  for  only  about  half  their  length.  Vesica  unusually  broad. 

Pubescence  suberect,  short;  tubercles  of  head  and  pronotum  bearing  longer,  erect  hairs  shorter  than 
three-quarters  of  the  diameter  of  an  eye. 

Colour  ochreous  yellow,  often  sprinkled  with  minute  red  dots.  Granules  of  antennae,  femora  and 
sometimes  of  tibiae  piceous;  antennal  segment  IV  often  piceous;  anterior  midline  of  pronotum,  apical  and 
basal  annuli  of  tibiae,  apical  halves  of  posterior  femora  and  more  or  less  extensive  markings  of 
connexivum,  especially  posterior  halves  of  laterotergites  IV- VI,  of  pleura,  of  thoracic  and  abdominal 
sterna  and  of  corium,  especially  in  two  places  on  the  apical  margin,  piceous. 

REMARKS.  The  short,  broad  vesica  and  only  partially  sclerotized  distal  dorsolateral  conjunctival  lobes  of 
this  species  are  unique  for  the  genus.  The  lack  of  ventral  sclerotized  conjunctival  strips,  the  small  body  size 
and  the  possession  of  an  antennal  segment  II  that  is  shorter  than  segment  III  are  all  shared  with  P.  parva. 
The  two  species  can  be  distinguished  externally  by  the  form  of  the  posterolateral  angles  of  the  abdominal 
sternites,  which  are  acute  in  parva  but  right-angled  in  brevitibialis . 


36 


37 


Figs  36-43  Psilolomia  species.  36,  37,  dispar:  (36)  dorsal  view  of  head;  (37)  lateral  view  of  head.  38,  39, 
vulgaris:  (38)  left  metathoracic  scent-gland  aperture;  (39)  metathoracic  wing.  40-43,  ventral  view  of 
abdominal  margin  of  (40)  steeleae;  (41)  lata;  (42)  brunneofusca;  (43)  vulgaris. 


174  W.  R.   DOLLING 

DISTRIBUTION.  Southern  India  and  Sri  Lanka. 

MATERIAL  EXAMINED 

Sri  Lanka:  1  $  (lectotype  of  brevitibialis),  Trincomalee  (Horn)  (IP).  India:  1  cf  (lectotype  of  typica), 
Chikkaballapura  (T.  V.  Campbell)  (BMNH). 

India:  26  cf ,  31  $,  data  as  holotype  of  N.  typica  (paralectotypes  of  typica)  (BMNH);  1  $,  Coimbatore, 
x.1953  (P.  5.  Nathan)  (IRSNB).  Sri  Lanka:  1  cf ,  Pundaloya  (BMNH). 

Psilolomia  parva  sp.  n. 

(Figs  31,  45,  51) 

Length:  cf ,  5-4-5-9  mm;  $,5-9-6-5  mm. 

Head  dorsally  granulate,  tuberculate  on  vertex  and  in  midline.  Antennifers  distinctly  but  rather  weakly 
divergent.  Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes  about  0-86.  Ratio  of 
lengths  of  antennal  segments  about  1-00:0-73:0-89:0-81.  Ratio  of  lengths  of  rostral  segments  about 
l-00:0-55:0-55:0-55.  Head  and  pronotum  together  constituting  more  than  0-39  (male)  or  0-35  (female)  of 
total  length. 

Pronotum  with  lateral  margins  (Fig.  31)  weakly  concave,  posterolateral  angles  scarcely  prominent; 
posterolateral  spines  acute,  at  least  1-5  times  as  long  as  their  basal  width;  width  of  pronotum  across  apices 
of  spines  divided  by  width  of  head  including  eyes  1-74-1-83.  Metathoracic  wing  with  antevannal  vein  weak, 
difficult  to  see.  Metasternum  rather  strongly  convex.  Anterior  and  intermediate  femora  without  or  with  a 
single  subapical  spine  beneath;  posterior  femur  with  three  major  spines,  the  first  about  half  as  long  as  the 
other  two,  one  to  three  minor  spines  between  the  two  large  spines  and  an  apical  series  of  four.  Length  of 
posterior  tibia  divided  by  length  of  posterior  femur  about  0-72. 

Abdominal  sternites  III- VII  with  posterolateral  angles  acute,  slightly  but  distinctly  prominent.  Male 
conjunctiva  (Fig.  45)  without  ventral  sclerotized  strips,  distal  dorsolateral  lobes  (Fig.  51)  short,  supported 
by  sclerotization  throughout  their  length;  vesica  slender. 

Pubescence  short,  semidecumbent  on  hemelytra  and  most  of  body,  slightly  longer  and  less  decumbent 
on  legs  and  antennae,  tubercles  of  head  and  pronotum  bearing  longer,  almost  erect  hairs  about  as  long  as 
two-thirds  diameter  of  an  eye. 

Colour  ochreous  yellow;  ground  colour  of  clavus,  corium  and  posterior  half  of  pronotum  pale  cinereous, 
without  yellow  tinge.  Grey  or  piceous  markings  of  greater  or  lesser  extent  present  on  midline  and  posterior 
border  of  pronotum,  on  scutellum,  on  hemelytra,  especially  costal  and  apical  areas  of  corium;  on  anterior 
and  posterior  borders  of  laterotergites,  including  more  than  posterior  halves  of  laterotergites  IV  and  V;  on 
apical  halves  of  femora,  especially  posterior  pair;  and  on  thoracic  pleura  and  abdominal  sterna.  Mesoster- 
num  and  metasternum  entirely  piceous.  Tibiae  with  basal,  apical  and  median  annuli  grey. 

REMARKS.  This  is  the  smallest  species  of  the  genus.  It  differs  from  the  African  P.  amphrysia  in  having  the 
second  antennal  segment  shorter  than  the  third  and  from  the  Asian  P.  brevitibialis  in  having  the 
posterolateral  angles  of  the  abdominal  sternites  acute.  The  antevannal  vein  is  very  indistinct,  probably 
because  it  is  functionally  obsolete  in  such  a  small  wing. 

DISTRIBUTION.  South-east  Asia. 

MATERIAL  EXAMINED 

Holotype  cf ,  Laos:  Wapikhamthong  Prov.,  Khong  Sedone,  15.viii.1965  (native  collector)  (BPBM). 

Paratypes.  Laos:  50  cf ,  49  $,  Wapikhamthong  Prov.,  Khong  Sedone,  various  dates  from  30. iv.  1965  to 
5.xi.l965  (native  collector);  13  Cf,  17  $,  Khong  Sedone,  3.viii.l965  (as  3.8.65)  (Rondon)  (BPBM; 
BMNH).  Thailand:  2  cf,  Chiangmai  Prov.,  Chiangmai  (Zoo),  16.vi.1965  (P.  D.  Ashlock);  3  cf,l  $, 
Chiangmai  Prov.,  Chiangmai  (Arboretum),  16.vi.1965  (P.  D.  Ashlock)  (BPBM). 

Psilolomia  dispar  (Walker)  comb.  n. 
(Figs  36,  37,  48) 

Clavigralla  dispar  Walker,  1872:  6.  Holotype  cf  (no  locality  cited)  (BMNH)  [examined]. 
[Clavigralla  aliena  (Walker);  Distant,  1901:  426.  Misidentification.] 

Length:  cf,  8-4-9-0  mm;  $,8-9-9-9 mm. 

Head  (Figs  36,  37)  dorsally  granulate,  with  antennifers  very  short,  slightly  divergent,  outer  apical 
process  obsolete;  a  large,  granular  tubercle,  almost  as  large  as  the  eye,  present  on  side  of  head  above  each 
antennifer.  Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes  0-81-0-90.  Ratio  of 


TRIBE  PSEUDOPHLOEINI  175 

lengths  of  antennal  segments  in  male  about  l-00:l-01:0-88:0-87,  in  female  about  l-00:0-99:0-84:0-92.  Ratio 
of  lengths  of  rostral  segments  about  1-00:0-63:0-55:0-61. 

Pronotum  with  lateral  margins  shallowly  concave,  granulate,  posterolateral  angles  elevated  but  only 
weakly  produced  laterally,  posterolateral  spines  acute,  about  1-5  times  as  long  as  their  basal  width;  width  of 
pronotum  across  apices  of  spines  divided  by  width  of  head  including  eyes  1-96-2-04.  Metasternum  weakly 
convex.  Anterior  femur  with  one  small  subapical  pine  beneath,  intermediate  femur  with  one  small  spine 
and  one  very  small  one,  posterior  femur  with  three  major  spines,  the  first  pair  with  no  minor  spines 
between  them,  the  second  and  last  with  three  or  four  minor  spines  between  them  and  with  an  apical  series 
of  four;  holotype  and  some  other  specimens  with  a  small  spine  preceding  the  three  major  spines. 

Abdominal  sternites  IH-VII  with  posterolateral  angles  slightly  more  acute  than  right-angled  and  slightly 
prominent.  Conjunctiva  (Fig.  48)  with  ventral,  toothed  sclerotized  strips  well  developed,  distal  dorso- 
lateral  lobes  sclerotized  throughout,  L-shaped. 

Coloration  generally  ferruginous;  dorsal  midline  of  head  with  distinct,  narrow,  yellow  stripe;  antennal 
segments  III-VII  yellow  with  granulation  piceous;  segment  IV  ferruginous;  pronotum  with  anteromedian 
macula  and  posterolateral  spines  brown;  legs  yellow  but  extensively  covered  with  brown  granules,  tibiae 
with  basal,  apical  and  median  annuli  brown,  femora  extensively  mottled  brown  or  ferruginous,  especially 
apically;  thoracic  sterna  brown;  abdominal  sterna  yellow  with  brown  markings  largely  confined  to  two 
longitudinal  bands;  laterotergites  yellow  with  brown  markings  medially  and  posteriorly;  clavus  and  corium 
ferruginous,  the  latter  with  a  few  obscure  spots  on  veins  brown,  membrane  fuscous,  veins  spotted  darker, 
apex  of  corium  and  membrane  throughout  with  large,  cream  spots. 

Pubescence  rather  short,  semidecumbent  to  suberect;  erect  hairs  about  as  long  as  two-thirds  diameter  of 
eye  present  rather  densely  on  declivent  part  of  pronotum  and  on  ventral  surfaces  of  femora;  main 
pubescence  of  head  consisting  of  short,  almost  decumbent  hairs  but  larger  granules,  especially  of  dorsal 
midline,  bearing  short,  suberect  to  erect  hairs. 

REMARKS.  This  species  is  immediately  recognizable  by  the  presence  of  the  large  tubercles  above  the 
antnnifers;  these  could  be  mistaken  for  the  antennifers  themselves  in  dorsal  view.  The  first  antennal 
segment  is  unusually  short  for  one  of  the  larger  species  of  the  genus.  Walker's  original  description  refers  to 
three  longitudinal  brown  bands  on  the  pronotum;  the  lateral  ones  are,  in  fact,  shadows  in  the  shallow 
troughs  between  the  gently  convex  pronotal  disc  and  the  elevated  posterolateral  angles.  Distant  (1901: 
426)  redescribed  the  type-specimen  of  Clavigralla  dispar  Walker  under  the  impression  that  it  was  the  type 
of  Cletus  alienus  Walker.  The  latter,  which  is  now  missing,  was  a  female  from  New  Guinea  whereas  the  type 
of  dispar  was  specified  by  Walker  as  a  male  without  locality  data  donated  by  Saunders. 

.DISTRIBUTION.  India. 

MATERIAL  EXAMINED 

1  cf  (holotype),  no  data  (Saunders)  (BMNH). 

India:  1  $ ,  Pondicherry  State,  Karikal,  vii.1963  (P.  S.  Nathan)  (BMNH);  2  cf ,  1  $  'E.  Ind.'  (Bacon);  1 
Cf ,  'Bgl'  (Haw.);  1  $,  no  locality  (Haw.)  (all  UM);  1  cf ,  Pondicherry  State,  Karikal,  l.i.1959  (Nathan);  1 
Cf,  Karikal  Terr.,  Kurumbaragum,  viii.1953  (Nathan);  6  cf ,  2  $ ,  Tranquebar,  vii.1953  (Nathan)  (J.  A. 
Slater  coll.). 

Psilolomia  vulgaris  sp.  n. 

(Figs  32,  38,  39,  43,  55) 

Length:  cf,  7-9-8-8  mm;  $,  8-7-9-7 mm. 

Head  dorsally  granulate.  Antennifers  abruptly  divergent.  Length  of  antennal  segment  I  divided  by  width 
of  head  including  eyes  in  male  about  0-89,  in  female  about  0-95.  Ratio  of  lengths  of  antennal  segments  in 
male  about  l-00:0-89:0-75:0-89,  in  female  about  l-00:0-96:0-81:0-93.  Ratio  of  lengths  of  rostral  segments 
about  l-00:0-56:0-56:0-56. 

Pronotum  (Fig.  32)  with  lateral  margins  very  shallowly  concave,  bearing  large,  conical  tubercles  about  as 
high  as  wide;  posterolateral  spines  short,  blunt,  not  longer  than  wide;  width  across  apices  of  spines  divided 
by  width  of  head  including  eyes  1-98-2-11.  Metasternum  moderately  convex.  Anterior  and  intermediate 
femora  each  without  or  with  a  single,  small  subapical  spine  beneath;  posterior  femur  with  three  major 
spines,  rarely  preceded  by  an  additional  small  spine,  the  first  pair  of  large  spines  without  or  with  a  single 
minor  spine  between  them,  three  or  four  minor  spines  between  the  middle  and  distal  spines  and  an  apical 
series  of  four.  Length  of  posterior  tibia  divided  by  length  of  posterior  femur  0-75-0-79.  Metathoracic  wing 
(Fig.  38)  and  peritreme  (Fig.  39)  typical  of  the  genus. 

Abdominal  sternites  III-VII  with  posterolateral  angles  slightly  more  acute  than  right-angled,  very 


176 


44 


W.  R.  DOLLING 


55 


Figs  44-57  Psilolomia  species.  44-46,  dorsal  view  of  conjunctiva  and  vesica  of  (44)  brevitibialis;  (45) 
parva;  (46)  brunneofusca.  47,  48,  ventral  view  of  conjunctiva  and  vesica  of  (47)  brunneofusca;  (48) 
dispar.  49-57,  apical  view  of  vesica,  ejaculatory  reservoir  apparatus  and  distal  dorsolateral  lobes  of 
conjunctiva  of  (49)  amphrysius;  (50)  brevitibialis;  (51) parva;  (52)  brunneofusca;  (53)  /ate;  (54)  lamottei; 
(55)  vulgaris;  (56)  pundaloyae;  (57)  nigeriensis. 


weakly  prominent  (Fig.  43).  Male  conjunctiva  with  ventral  sclerotized  strips  rather  weakly  developed, 
distal  dorsolateral  lobes  fully  sclerotized,  strongly  angled  before  their  spatulately  expanded  apices  (Fig. 

Antennal  segments  I-III  with  suberect  pubescence  shorter  than  width  of  segment;  rostrum  with  short, 
suberect  and  decumbent  pubescence;  head  with  short,  semidecumbent  hairs  and  dorsally  with  longer, 
suberect  hairs  not  confined  to  midline;  pronotum  with  short,  semidecumbent  pubescence  posteriorly  and 
suberect  pubescence  on  disc  and  anteriorly,  interspersed  except  posteriorly  with  rather  longer,  erect  hairs; 
legs  with  short  to  moderately  long,  suberect  pubescence,  femora  with  longer,  erect  hairs  ventrally;  clavus 
and  corium  with  short,  semidecumbent  pubescence;  thoracic  pleura  and  sterna  and  abdominal  sterna  and 
laterotergites  with  slightly  longer,  semidecumbent  pubescence. 

Coloration  generally  pale  yellowish  brown;  ocellar  tubercles  and  granules  of  antennae,  rostrum  and  legs 
brown.  Pronotum  with  anteromedian  macula,  lateral  granules  and  posterolateral  spines  in  part  brown; 
scattered  markings  on  pleura,  scutellum  and  corium,  especially  apical  areas  of  corium,  brown;  thoracic 
sterna  piceous;  femora  sparsely  mottled  with  brown,  apical  half  of  posterior  femur,  especially  on  posterior 
face,  reddish  brown.  Abdominal  laterotergites  and  bands  along  abdominal  sterna  mottled  dark  brown. 
Membrane  of  hemelytra  very  palely  infuscate,  veins  paler  with  interrupted  brown  streaks. 


TRIBE  PSEUDOPHLOEINI  177 

REMARKS.  This  species  closely  resembles  P.  pundaloyae  and  the  African  P.  nigeriensis;  it  differs  from  the 
former  in  the  shorter  pronotal  posterolateral  spines,  from  the  latter  in  the  longer  pubescence  of  the  head 
and  from  both  in  the  form  of  the  distal  dorsolateral  lobes  of  the  conjunctiva.  The  sexual  dimorphism  in  the 
relative  length  of  antennal  segment  I  is  unusual;  the  shortness  of  this  segment  in  the  female  results  in  the 
length  of  segment  IV  divided  by  that  of  segment  I  being  greater  in  the  female  than  in  the  male,  the  reverse 
of  the  usual  situation  in  the  subfamily,  where  segment  IV  is  relatively  longer  in  the  male. 

DISTRIBUTION.  Southern  India,  Sri  Lanka. 

MATERIAL  EXAMINED 

Holotype  cf ,  India:  Madras,  Karikal,  Kurumbagarum,  xi.1953  (P.  S.  Nathan)  (J.  A.  Slater  coll.). 

Paratypes.  India:  8  Cf,  1  $,  Karikal  Territory,  Kurumbagarum,  ix.1954  (Nathan)  (IRSNB);  1  cf, 
Karikal,  v.1957  (Nathan);  1  cf,  Tranquebar,  viii.1956  (Nathan)  (BPBM);  1  cf,  2  $,  Pondicherry  or 
Karikal,  vii.1953  (Nathan)  (BMNH);  9  specimens,  Tranquebar,  vii.1953  (Nathan);  36  specimens,  Kurum- 
bagarum, various  dates  vii-xi.1953  and  viii.1954  (Nathan)  (J.  A.  Slater  coll.). 

Psilolomiapundaloyaesp.  n. 

(Figs  1,56) 

Length:  cf ,  8-2-8-8  mm;  $,9-1  mm. 

Head  dorsally  granulate.  Antennifers  rather  strongly  divergent.  Length  of  antennal  segment  I  divided  by 
width  of  head  including  eyes  1-00-1-05.  Ratio  of  lengths  of  antennal  segments  about  1-00:0-92:0-75:0-80 
(IV  missing  in  female).  Ratio  of  lengths  of  rostral  segments  about  l-00:0-58:0-49:0-56. 

Pronotum  (Fig.  1)  with  lateral  margins  shallowly  concave,  bearing  short  tubercles  slightly  longer  than 
their  basal  width;  posterolateral  spines  longer  than  wide;  width  across  apices  of  spines  divided  by  width  of 
head  including  eyes  2-00-2-12.  Metasternum  weakly  convex,  its  anterior  emargination  extending  post- 
eriorly as  a  shallow  sulcus  for  over  half  its  length.  Anterior  femur  with  a  minute  subapical  spine  beneath, 
intermediate  femur  with  none,  posterior  femur  with  three  major  subapical  spines,  one  minor  spine 
between  the  first  two,  three  between  second  and  third  and  an  apical  series  of  four.  Length  of  posterior  tibia 
divided  by  that  of  posterior  femur  about  0-82. 

Abdominal  sternites  (Fig.  1)  with  posterolateral  angles  distinctly  acute  and  slightly  prominent.  Male 
conjunctiva  with  ventral,  toothed,  sclerotized  strips  well  developed;  distal  dorsolateral  lobes  (Fig.  56) 
wholly  sclerotized,  not  angled  before  the  spatulate  apex. 

Pubescence  rather  short,  suberect,  tending  to  semidecumbent  on  ventral  surface  of  body;  longer,  erect 
hairs  throughout  declivent  part  of  pronotum  and  dorsal  surface  of  head.  Longest  hairs  of  head  and 
pronotum  more  than  two-thirds  as  long  as  diameter  of  an  eye. 

Coloration  pale  yellowish  brown  with  darker  markings,  as  in  P.  vulgaris. 

REMARKS.  This  species  closely  resembles  P.  vulgaris  but  differs  in  the  longer  and  more  slender  postero- 
lateral pronotal  spines  and  the  longer  pubescence. 

DISTRIBUTION.  India,  Sri  Lanka. 

MATERIAL  EXAMINED 

Holotype  $,  Sri  Lanka:  Pundaloya  (Green)  (BMNH). 

Paratypes.  India:  1  cf,  Chikkaballapura,  i.1915  (Campbell);  1  cf,  Kodai  Kanal,  v.1918  (Campbell) 
(BMNH). 

Psilolomia  brunneofusca  sp.  n. 

(Figs  33, 42,  46,  47,  52) 

Length:  cf ,  7-6-8-8  mm;  $,7-9-8-5  mm. 

Head  dorsally  granulate.  Antennifers  abruptly  divergent.  Length  of  antennal  segment  I  divided  by  width 
of  head  including  eyes  1-15-1-20.  Ratio  of  lengths  of  antennal  segments  in  male  about  1 -00:0-81 :0-68:0-74, 
in  female  about  l-00:0-83:0-77:0-73.  Ratio  of  lengths  of  rostral  segments  about  1-00:0-62:0-56:0-60. 

Lateral  margins  of  pronotum  (Fig.  33)  concave,  bearing  granules  about  as  high  as  wide;  posterolateral 
spines  acute,  longer  than  their  basal  width  and  slightly  curved  posteriad;  width  across  apices  of  spines 
divided  by  width  of  head  including  eyes  2-11-2-14.  Metasternum  weakly  convex.  Anterior  femur  with  a 
small,  subapical  spine  beneath,  sometimes  with  an  incipient  second  one  proximal  to  it;  intermediate  femur 
with  two  spines;  posterior  femur  with  three  major  spines  and  usually  a  fourth,  smaller  one  proximal  to  the 
others,  one  to  three  minor  spines  between  the  first  pair  of  large  spines,  three  to  four  between  the 


178  W.  R.  DOLLING 

penultimate  and  last  and  an  apical  series  of  four  or  five.  Length  of  posterior  tibia  divided  by  that  of 
posterior  femur  0-78-0-80. 

Abdominal  sternites  III-VII  (Fig.  42)  with  posterolateral  angles  acute  and  prominent.  Male  conjunctiva 
(Figs  46,  47)  with  ventral  sclerotized  strips  well  developed  and  distal  dorsolateral  lobes  (Fig.  52)  fully 
sclerotized,  with  slightly  expanded  apices. 

Pubescence  short,  semidecumbent;  rather  longer  and  suberect  on  legs,  antennae,  declivent  part  of 
pronotum  and  dorsum  of  head ;  length  of  hairs  of  suberect  pubescence  equalling  half  to  two-thirds  diameter 
of  an  eye. 

General  colour  dark  ochreous  brown  to  almost  black.  Antennae  and  legs  yellow  with  all  granulation 
dark  brown,  antennal  segment  IV  strongly  infuscate,  apical  halves  of  anterior  and  intermediate  tibiae 
mottled  with  dark  brown,  apical  half  of  posterior  femur  heavily  mottled  dark  brown.  Head,  pronotum  and 
scutellum  ochreous  with  dark  brown  markings,  the  raised,  white  apex  of  the  scutellum  (universal  in  the 
genus)  strongly  contrasting.  Thoracic  pleura  and  abdominal  sterna  and  laterotergites  reddish  yellow  with 
more  or  less  extensive  dark  brown  markings;  thoracic  sterna  piceous  or  black.  Clavus  and  corium  dark 
ochreous  or  grey-brown,  main  veins  of  corium  spotted  both  darker  and  paler;  membrane  darkly  infuscate, 
veins  brown;  membrane  and  apex  of  corium  with  more  or  less  conspicuous  whitish  spots,  those  of 
membrane  often  interrupting  darker  colour  of  veins. 

REMARKS.  This  is  a  widespread,  dark  brown  species  similar  in  build  to  the  much  paler,  yellowish  P.  vulgaris. 
It  may  be  synonymous  with  P.  davipes,  from  southern  China,  but  specimens  of  the  latter  were  not  available 
for  study.  It  may  be  that  the  abdominal  sternites  of  davipes  lack  the  acutely  produced  posterolateral  angles 
that  are  more  developed  in  P.  brunneofusca  than  in  any  other  species  of  the  genus.  The  Indian  specimen 
examined  differed  from  the  south-east  Asian  specimens  only  in  having  slightly  broader  posterolateral 
pronotal  spines.  The  specimen  from  peninsular  Malaya  reported  below  represents  the  most  south-easterly 
record  of  any  species  of  the  tribe;  only  some  Clavigrallini  represent  the  subfamily  in  the  Malay  Archipelago 
and  northern  Australia. 

DISTRIBUTION.  South-eastern  India,  Laos,  Malaya. 

MATERIAL  EXAMINED 

Holotype  cf ,  Laos:  Pakse,  iii.1965  (/.  A.  Rondori)  (BPBM). 

Paratypes.  Laos:  1  $,  Wapikhamthong  Prov.,  Khong  Sedone,  18. ix. 1965  (native collector)  (BMNH);  1 
Cf,  Borikhame  Prov.,  Pakkading,  100-200  m,  23. iv.  1965  (J.  L.  Gressitt);  1  cf,  Vientiane  Prov.,  Tha 
Ngone,  3.i.l965  (native  collector)  (BPBM);  1  cf ,  Luang  Prabang,  Pak  Leung,  5.iii.l920  (R.  V.  de  Salvaza) 
(BMNH).  India:  1  cf,  Anamalai  Hills,  Cinchona,  3500  ft  [1070  m],  v.1957  (P.  S.  Nathan)  (BMNH). 
Malaya:  1  cf ,  Perlis,  Kaki  Bukit,  27.V.74  (P.  Roche)  (BMNH). 

Psilolomia  davipes  (Hsiao)  comb.  n. 

Psilocoris  davipes  Hsiao,  1964: 251-252;  259.  Holotype  $ ,  CHINA:  Yunnan,  1250  m,  17.vi.1956  (Academia 
Sinica,  Beijing)  [not  examined]. 

Length:  cf ,  unknown;  $,8-7  mm. 

Length  of  antennal  segment  I  about  1  •  15  times  width  of  head  including  eyes.  Ratio  of  lengths  of  antennal 
segments  as  1-00:0-87:0-80:0-80.  Ratio  of  lengths  of  rostral  segments  as  1-00:0-69:0-54:0-61.  Pronotum 
with  posterolateral  angles  not  produced;  posterolateral  spines  longer  than  their  basal  width. 

General  coloration  'dark  ochraceus'. 

REMARKS.  The  partial  description  given  here  is  based  on  that  of  Hsiao  (1964:  251-252,  259).  No  material 
was  available  for  study.  A  photograph  of  this  species  is  reproduced  by  Hsiao  (1977:  pi.  46,  fig.  603).  It  looks 
very  similar  to  P.  brunneofusca  and  the  two  species  may  be  synonymous. 

DISTRIBUTION.  Southern  China  (Yunnan). 

Psilolomia  steeleae  sp.  n. 

(Figs  34,  40) 

Length:  cf ,  unknown;  $ ,  8-3  mm. 

Head  dorsally  granulate.  Antennifers  weakly  divergent.  Length  of  antennal  segmentl  divided  by  width 
of  head  including  eyes  1-03.  Ratio  of  lengths  of  antennal  segments  as  l-00:0-86:0-76:0-79.  Ratio  of  lengths 
of  rostral  segments  as  l-00:0-64:0-54:0-68. 

Pronotum  (Fig.  34)  with  lateral  margin  distinctly  concave,  bearing  tubercles  slightly  longer  than  their 


TRIBE  PSEUDOPHLOEINI  179 

basal  width;  posterolateral  spines  short,  broadly  triangular;  width  across  apices  of  spines  divided  by  width 
of  head  including  eyes  1  -91 .  Metasternum  almost  flat.  Anterior  and  intermediate  femora  each  with  a  small, 
subapical  spine  beneath;  posterior  femur  with  three  major  spines,  the  first  half  as  long  as  the  other  two,  no 
minor  spines  between  this  and  the  middle  spine,  two  minor  spines  between  the  two  large  spines  and  an 
apical  series  of  three.  Posterior  tibiae  rather  long,  0-88  times  as  long  as  posterior  femur. 

Abdominal  sternites  III-VII  with  posterolateral  angles  (Fig.  40)  slightly  more  acute  than  right-angled, 
weakly  prominent. 

Pubescence  short,  semidecumbent;  rather  longer  and  suberect  on  legs,  antennae,  declivent  part  of 
pronotum  and  dorsum  of  head;  length  of  hairs  of  suberect  pubescence  equalling  half  to  two-thirds  diameter 
of  an  eye. 

Coloration  paler  than  in  P.  brunneofusca  but  not  as  pale  as  in  P.  vulgaris;  darker  markings  distributed  as 
in  vulgaris;  membrane  of  hemelytra  fuscous  with  traces  of  pale  spotting. 

REMARKS.  This  new  species  differs  from  P.  vulgaris  in  the  longer  first  antennal  segment  and  darker  colour, 
and  from  P.  brunneofusca  in  its  shorter  pronotal  and  abdominal  spines. 

DISTRIBUTION.  Known  only  from  the  type-locality  in  Burma. 

MATERIAL  EXAMINED 
Holotype  $,  Burma:  Mishmi  Hills,  Lohit  River,  24.iii.1935  (M.  Steele)  (BMNH). 

Psilolomia  lata  sp.  n. 

(Figs  35,  41,  53) 

Length:  d",  7-8-8-5  mm;  $,8-5-9-3  mm. 

Head  dorsally  granulate.  Antennifers  rather  strongly  and  often  abruptly  divergent.  Length  of  antennal 
segment  I  divided  by  width  of  head  including  eyes  1-03-1  •  16.  Ratio  of  lengths  of  antennal  segments  in  male 
about  l-00:0-90:0-76:0-79,  in  female  about  1-00:0-90:0-77:0-70.  Ratio  of  lengths  of  rostral  segments  about 
1-00:0-67:0-53:0-60. 

Pronotum  (Fig.  35)  with  lateral  margins  deeply  concave,  bearing  tubercles  slightly  longer  than  their 
basal  width;  posterolateral  spines  short,  triangular;  width  across  apices  of  spines  divided  by  width  of  head 
including  eyes  1-96-2-09.  Metasternum  flat,  midline  broadly  and  very  shallowly  concave  throughout. 
Anterior  and  intermediate  femora  each  bearing  a  single  subapical  spine,  rarely  a  second  spine  present  on 
intermediate  femur;  posterior  femur  with  three,  rarely  four,  major  spines,  penultimate  spine  preceded  by  a 
single  minor  spine  or  not,  succeeded  by  three  or  four  minor  spines;  terminal  series  consisting  of  four,  rarely 
five,  spines.  Posterior  tibia  rather  long,  its  length  divided  by  that  of  posterior  femur  0-86-0-87. 

Abdominal  sternites  III-VII  with  posterolateral  angles  (Fig.  41)  slightly  more  acute  than  right-angled, 
not  or  very  slightly  prominent.  Conjunctiva  with  ventral  sclerotized  strips  well  developed;  distal  dorso- 
lateral  lobes  (Fig.  53)  supported  by  sclerotization  right  up  to  their  apices,  which  are  not  expanded. 

Pubescence  short,  semidecumbent;  rather  longer  and  suberect  on  legs,  antennae,  declivent  part  of 
pronotum  and  dorsal  midline  of  head;  length  of  hairs  of  suberect  pubescence  equalling  half  to  two-thirds 
diameter  of  an  eye. 

General  coloration  ferruginous,  dark  markings  distributed  as  in  P.  vulgaris;  membrane  of  hemelytra 
slightly  infuscate,  veins  brown,  interrupted  by  paler  stretches. 

REMARKS.  This  broad-bodied,  ferruginous  species  may  be  distinguished  from  all  others  of  the  genus  by  the 
very  deeply  concave  pronotal  lateral  margins. 

DISTRIBUTION.  Thailand. 

MATERIAL  EXAMINED 

Holotype  C?,  Thailand:  Chiangmai  Prov.,  Doi  Pui,  17.vi.1965  (P.  D.  Ashlock)  (BPBM). 

Paratypes.  Thailand:  4  cf,  data  as  holotype;  2  d",  2  $,  Chiangmai,  Doi  Suthep,  1300  m,  8.vi.l965 
(Ashlock)  (BPBM;  BMNH). 

PUNGRA  gen.  n. 

Type-species:  Pungra  angusta  sp.  n. 

Body  form  rather  narrowly  oblong,  slightly  depressed,  connexivum  slightly  widened. 

Head  longer  than  pronotum.  Antennifers  moderately  divergent,  outer  apical  processes  porrect,  broadly 
triangular.  Antennal  segment  I  about  as  long  as  head  width  across  antennifers,  abruptly  but  weakly 


180  W.  R.  DOLLING 

clavate,  slightly  longer  than  II,  distinctly  shorter  than  III,  IV  equal  to  I  in  male,  shorter  in  female.  Bucculae 
occupying  about  one-quarter  of  length  of  ventral  midline  of  head.  Rostrum  at  rest  reaching  to  anterior  disc 
of  metasternum,  segment  II  subequal  to  I,  IV  much  shorter,  III  shortest. 

Pronotum  gently  declivent,  much  broader  than  long,  with  a  pair  of  small,  raised  calli  on  posterior  disc 
between  posterolateral  angles,  posterolateral  angles  weakly  produced,  posterolateral  spines  small,  lateral 
margins  with  four  stout  tubercles  each,  posterior  margin  weakly  convex,  prescutellar  spines  absent. 
Scutellum  almost  flat,  equilateral,  its  apex  scarcely  raised.  Mesosternum  and  metasternum  deeply  sulcate 
throughout.  Dorsal  ridge  of  metathoracic  peritreme  reniform.  Metathoracic  wing  with  antevannal  vein. 
Anterior  and  intermediate  femora  without  spines  or  tubercles,  posterior  femur  with  basal  tubercle  well 
developed,  with  two  major  subapical  spines  preceded  by  granules  and  with  three  or  four  spine-like 
tubercles  between  them  and  an  apical  series  of  four  tubercles. 

Abdominal  sternites  with  posterolateral  angles  only  slightly  prominent,  lateral  margins  not  tuberculate. 
Male  genital  capsule  emarginate,  emargination  filled  by  apices  of  parameres. 

REMARKS.  This  rather  undistinguished  genus  is  probably  close  to  Pseudomyla,  which  it  resembles  in  the 
presence  of  stout  tubercles  on  the  lateral  margins  of  the  pronotum,  but  differs  from  it  in  the  shape  of  the 
posterolateral  pronotal  angles. 

DISTRIBUTION.  India. 

Pungra  angusta  sp.  n. 

(Fig.  58) 

Length:  cf,  7-3-7-7 mm;  $,8-0-8-4 mm. 

Length  of  antennal  segment  I  0-76-0-89  times  width  of  head  across  eyes;  ratio  of  lengths  of  segments 
about  1-00:0-96:1-28:1-00  (male)  or  l-00:0-98:l-32:0-85  (female);  specialized  sensory  area  of  segment  IV 
occupying  about  five-sixths  of  its  length.  Ratio  of  lengths  of  rostral  segments  about  l-00:0-92:0-30:0-56. 

Pronotum  (Fig.  58)  with  lateral  margins  straight  for  much  of  their  length,  posterolateral  angles  abruptly 
but  shortly  produced,  posterolateral  spines  short,  width  across  apices  of  spines  1-63-1-68  times  width  of 
head  across  eyes.  Calli  of  posterior  disc  granular,  close  together.  Length  of  posterior  tibia  0-84-0-86  times 
length  of  posterior  femur. 

Abdominal  sternites  with  posterolateral  angles  acute,  very  slightly  prominent.  Aedeagus  similar  in  all 
respects,  including  the  unpaired  apical  ventral  lobe,  to  that  of  Pseudomyla  spinicollis  except  that  distal 
dorsomedian  lobe  is  broadly  conical  and  lacks  finger-like  processes  at  the  lateral  angles. 

Antennal  segment  I  strongly,  II  and  III  weakly  granulate.  Head  granulate,  dorsally  with  a  few  larger 
granules  and  tubercles.  Pronotum  weakly  granulate,  densely  punctate,  with  a  few  tubercles  anteriorly  in 
addition  to  the  four  stout  tubercles  on  the  lateral  margins.  Scutellum,  abdominal  sternites  including  their 
lateral  margins  and  laterotergites  weakly  granulate,  thoracic  pleura  moderately  granulate-punctate, 
thoracic  sterna  weakly  punctate.  Clavus  and  corium  strongly,  seriately  punctate  except  for  impunctate  area 
near  middle  of  apical  margin  of  corium.  Femora  rather  weakly  granulate. 

Pubescence  of  antennae,  head,  thorax,  femora  and  abdomen  of  short,  crisped,  semidecumbent,  pale, 
rather  flattened  hairs.  Femora,  tibiae  and  tarsi  with  short,  suberect,  fine  hairs.  Tubercles  of  head  and 
pronotum  with  short,  erect,  curved,  flattened  hairs. 

Colour  yellowish  stramineous,  ocellar  tubercles  dark  brown,  antennal  segment  I,  sides  of  head, 
pronotum,  pleura,  thoracic  and  abdominal  sterna  with  diffuse  brown  markings;  apices  of  all  femora, 
especially  the  posterior  pair,  with  brown  mottling,  all  tibiae  with  proximal  and  distal  incomplete  brown 
annuli,  anterior  tibiae  also  with  incomplete  median  brown  annuli.  Membrane  of  hemelytra  colourless,  its 
veins  white,  veins  of  membrane  and  of  corium  with  a  few  brown  spots.  Laterotergites  with  small  brown 
patches  at  base,  at  apex  and  in  middle  of  lateral  margins. 

DISTRIBUTION.  India. 

MATERIAL  EXAMINED 
Holotype  C?,  India:  Nagpur  (BMNH). 
Paratypes.  India:  1  $,  Bombay;  1  CM  $,  Maharashtra,  Jalna,  4.ix.l971  (/.  C.  Deeming)  (BMNH). 

PSEUDOMYLA  gen.  n. 

Type-species:  Merocoris  spinicollis  Spinola. 

Body  rather  narrowly  oblong,  not  depressed,  connexivum  moderately  widened. 
Head  shorter  than  pronotum;  antennifers  moderately  divergent,  outer  apical  processes  of  antennifers 


TRIBE  PSEUDOPHLOEINI 


181 


58 


Figs  58-64  58,  Pungra  angusta,  dorsal  view  of  head  and  pronotum.  59-63,  Pseudomyla  spinicollis:  (59) 
dorsal  view  of  conjunctiva  and  vesica;  (60)  ventral  view  of  same;  (61)  left  lateral  view  of  same  (dorsal  to 
left  of  figure);  (62)  ventral  view  of  abdominal  margin;  (63)  dorsal  view  of  pronotum.  64,  P.  cornuta, 
dorsal  view  of  pronotum. 


about  as  long  as  broad,  porrect.  Antennal  segment  I  abruptly  but  weakly  clavate,  as  long  as  or  slightly 
shorter  than  head;  segments  I-IV  subequal  in  length;  specialized  sensory  area  occupying  about  eight- 
ninths  of  length  of  antennal  segment  IV.  Bucculae  occupying  about  one-quarter  of  ventral  midline  of  head. 
Rostrum  at  rest  reaching  to  anterior  margin  or  disc  of  metasternum,  segment  II  almost  as  long  as  I,  IV 
about  two-thirds  as  long  and  III  about  two-fifths  as  long  as  I. 

Pronotum  (Figs  63,  64)  with  posterolateral  angles  strongly  produced  and  directed  forwards,  strongly 
declivent,  posterior  margin  straight  or  very  shallowly  emarginate,  without  prescutellar  spines,  lateral 
margins  each  with  three  or  four  stout  tubercles,  disc  between  posterolateral  angles  with  a  pair  of  small, 
transverse,  granular  calli.  Scutellum  almost  flat  with  apex  and  anterior  angles  slightly  elevated.  Mesoster- 
num  and  metasternum  sulcate.  Dorsal  ridge  of  metathoracic  peritreme  reniform.  Metathoracic  wing  with 
antevannal  vein  present  but  feebly  developed.  Anterior  and  intermediate  femora  with  or  without  a  single 
subapical  spine  beneath;  posterior  femur  with  a  well-developed  basal  tubercle,  three  major  subapical 
spines  of  which  the  proximal  one  is  about  half  as  long  as  the  others,  two  or  three  tubercles  between  the  two 
largest  spines  and  an  apical  series  of  three  or  four  tubercles.  Posterior  tibia  0-8-0 -9  times  as  long  as  femur. 

Abdominal  sternites  III-VII  with  posterolateral  angles  produced  into  small  spines  (Fig.  62).  Male 
genital  capsule  posteriorly  emarginate,  emargination  filled  by  apices  of  parameres.  Spermatheca  with  bulb 
gibbously  lunate,  bulb  short. 

REMARKS.  These  reddish  ochreous  insects  with  strongly  produced  and  anteriorly  directed  posterolateral 
pronotal  angles  have  been  placed  in  the  African  genus  Myla  on  the  basis  of  the  wholly  superficial 
similarities  of  body  shape.  The  genitalia  of  the  males  are  quite  unlike  the  characteristic  form  encountered 
in  that  genus  and  the  nearest  relative  of  Pseudomyla  may  be  Pungra,  which  it  resembles  in  the  presence  of 
stout  tubercles  on  the  lateral  margins  of  the  pronotum  and  granular  calli  on  its  posterior  disc;  the  form  of 
the  conjunctiva  is  very  similar  in  the  two  genera. 

DISTRIBUTION.  Oriental  region. 

Key  to  species 

1  Pronotum  (Fig.  63)  with  posterolateral  margins  behind  the  posterolateral  spines  weakly 

concave,  almost  straight.  (Central  and  southern  India,  Sri  Lanka) spinicollis  (p.  182) 

-  Pronotum  (Fig.  64)  with  posterolateral  margins  behind  the  posterolateral  angles  distinctly 

convex.  (S.  China,  Laos) cornuta(p.  182) 


182  W.  R.  DOLLING 

Pseudomyla  spinicollis  (Spinola)  comb.  n. 
(Figs  59-63) 

Merocoris  spinicollis  Spinola,  1837:  216-217.  Syntype(s),  INDIA:  Bombay  (Duponf)  [lost]. 
Clavigralla  concolor  Dohrn,  1860:  403.  LECTOTYPE  cf,  SRI  LANKA  (IZPAN),  here  designated  [ex- 
amined]. Syn.  n. 
Myla  concolor  (Dohrn)  Stal,  1873:  84. 

Length:  cf ,  6-75-7-1  mm;  $  unknown. 

Length  of  antennal  segment  I  equal  to  width  of  head  including  eyes;  ratio  of  lengths  of  antennal  segments 
about  1  -00:0-92: 1  -00:0-96.  Ratio  of  lengths  of  rostral  segments  about  1  -00:0-91 :0-41 :0-68. 

Pronotum  (Fig.  63)  with  posterolateral  margins  (extending  backwards  from  posterolateral  spines)  very 
weakly  concave,  almost  straight.  Metasternum  moderately  convex,  shallowly  sulcate  throughout.  Anterior 
and  intermediate  femora  each  with  a  single,  small  (Sri  Lanka)  or  obsolete  (Bombay)  subapical  spine. 

Conjunctiva  (Figs  59-61)  with  M-shaped  dorsomedian  lobe  with  pronounced  dorsal  angles;  low,  broad 
distal  dorsomedian  lobe  with  two  anteriorly  projecting  finger-like  appendages;  distal  dorsolateral  lobes 
descending,  supported  throughout  their  ventral  faces  by  a  sclerotized  strip  articulating  with  apices  of  wings 
of  ejaculatory  reservoir  complex;  apical  ventral  lobe  single,  weakly  sclerotized;  distal  ventrolateral  lobes 
membranous  except  for  a  pair  of  toothed  strips  in  ventral  wall,  which  project  at  their  anterior  ends  on  very 
small,  free  lobes.  Vesica  slender,  protected  at  base  by  a  single,  asymmetrical  sclerite;  ejaculatory  reservoir 
asymmetrical  but  with  wings  symmetrical,  straps  absent. 

Granulation  of  antennal  segment  I  dense,  uniform,  that  of  segments  II  and  III  sparser  and  smaller;  head 
granulate,  pronotum  and  all  pleura  strongly  punctate  and  rather  weakly  granulate,  scutellum  moderately 
granulate-punctate,  thoracic  and  abdominal  sterna  weakly  granulate,  laterotergites  weakly  granulate-, 
punctate,  clavus  and  corium  densely  and  strongly  sedately  punctate  except  for  impunctate  area  near 
middle  of  apical  margin  of  corium,  femora  and  tibiae  sparsely  and  weakly  granulate. 

Semidecumbent,  short,  white  pubescence  of  flattened  hairs  present  on  antennal  segments  I-III,  head, 
thorax,  clavus,  corium  and  femora;  hairs  of  thoracic  pleura,  pronotum  and  scutellum  confused,  those  of 
other  areas  lying  more  uniformly  parallel;  femora,  tibiae  and  tarsi  with  rather  longer,  fine,  colourless, 
semi-erect  pubescence. 

Entire  insect  ochreous  red  except  for  black  meso-  and  metasterna,  diffuse  blackish  markings  on  ventral 
surfaces  of  head,  pronotum  and  abdomen  and  colourless  hemelytral  membrane  with  veins  marked  white 
and  brown;  tibiae  paler  red  than  rest  of  insect. 

REMARKS.  Spinola's  type-material  of  Merocoris  spinicollis,  sent  him  by  Dupont  from  Bombay,  could  not  be 
found  in  Spinola's  collection  by  Prof.  Dott.  Carlo  Vidano,  of  Turin,  who  kindly  searched  for  it  on  my 
behalf,  nor  is  it  catalogued  by  Casale  (1981).  The  genus  is  of  such  distinctive  appearance  that  there  can  be 
no  doubt  that  spinicollis  belongs  here.  Spinola's  description  is  clear  and  the  length  given  (3  French  lines: 
equal  to  7  mm)  is  within  the  measured  range  of  specimens  available.  The  slight  difference  in  the  size  of  the 
spines  of  the  anterior  and  intermediate  femora  of  the  specimens  from  Bombay  and  those  from  Sri  Lanka 
does  not  seem  sufficient  to  warrant  a  division  of  the  material  into  two  species,  particularly  in  view  of  the 
small  number  of  specimens  available.  If  it  should  prove  that  the  Sri  Lanka  specimens  are  specifically 
distinct  from  the  Bombay  ones,  Dohrn's  name  concolor  is  available  for  the  species. 

DISTRIBUTION.  India  and  Sri  Lanka.  Hsiao's  (1964:  252)  record  of  Myla  concolor  from  China  refers  to  the 
material  upon  which  he  later  (Hsiao,  1965:  427)  founded  the  following  species. 

MATERIAL  EXAMINED 

Sri  Lanka:  1  cf  (lectotype  of  concolor),  (Nietner)  (IZPAN). 

Sri  Lanka:  1  cf ,  Pundaloya  (BMNH);  1  cf ,  Pundaloya  (Atkinson)  (UM).  India:  2  cf ,  Bombay  (Dixon), 
ICf,  Bombay  (BMNH). 

Pseudomyla  cornuta  (Hsiao)  comb.  n. 
(Fig.  64) 

Myla  cornuta  Hsiao,   1965:  427,  433.  Holotype   $,  CHINA:  Yunnan,  Xishuang,  Banna,  20.V.1958 
(Academia  Sinica,  Beijing)  [not  examined]. 

Length:  cf ,  7-0  mm  (fide  Hsiao,  1965);  $ ,  6-5-7-2  mm. 

Length  of  antennal  segment  I  0-94  times  width  of  head  including  eyes;  ratio  of  lengths  of  antennal 
segments  about  l-00:0-90:l-06:0-88.  Ratio  of  lengths  of  rostral  segments  about  l-00:0-95:0-41:0-68. 


TRIBE  PSEUDOPHLOEINI  183 

Pronotal  margins  behind  posterolateral  spines  gently  but  distinctly  convex  (Fig.  64).  Metasternum 
moderately  convex,  rather  deeply  sulcate  throughout.  Anterior  and  intermediate  femora  unarmed  (fide 
Hsiao)  or  with  a  single,  small  subapical  spine  beneath.  Posterior  femur  with  three  major  spines,  the  most 
proximal  less  than  one-third  as  long  as  the  others,  between  which  are  two  to  four  tubercles  and  beyond 
which  is  an  apical  series  of  four  tubercles.  Coloration,  especially  of  clavus  and  corium,  more  ochreous  than 
that  of  P.  spinicollis. 

REMARKS.  Type-material  was  not  available  but  the  material  examined  agrees  rather  closely  with  the 
original  description,  and  the  locality  from  which  it  was  obtained  is  near  to  the  type-locality. 

DISTRIBUTION.  China:  Yunnan  (type-locality)  and  Laos. 

MATERIAL  EXAMINED 
Laos:  4  $,  Sedone  Province,  Paksong,  18.V.1965  (P.  D.  Ashlock)  (BPBM). 

NEOMEVANIOMORPHA  gen.  n. 

Type-species:  Mevaniomorpha  annulipes  rodhaini  Schouteden,  1938:  296. 

Body  oblong,  neither  compressed  nor  depressed,  connexivum  broad. 

Head  about  as  long  as  pronotum;  eyes  small,  prominent.  Antennifers  divergent,  outer  apical  process 
about  as  long  as  broad,  porrect.  Antennal  segments  subequal  in  length,  segment  I  weakly  clavate, 
granulate  to  tuberculate,  distinctly  shorter  than  head,  segments  II  and  III  slender,  very  weakly  granulate, 
IV  elongate  fusiform  with  specialized  sensory  area  occupying  about  seven-eighths  of  its  length.  Bucculae 
occupying  about  one-quarter  of  ventral  midline  of  head.  Rostrum  at  rest  reaching  disc  of  metasternum; 
segment  III  the  shortest,  II  and  IV  subequal,  slightly  shorter  than  I. 

Pronotum  (Fig.  65)  rather  strongly  declivent,  granulate-tuberculate,  lateral  margins  bearing  several 
short  tubercles,  posterolateral  angles  strongly  produced  anterolaterally,  tapering  to  terminal  spine; 
posterior  margin  weakly  convex,  with  a  few  granules,  prescutellar  spines  absent.  Scutellum  almost  flat, 
equilateral,  apex  not  elevated  or  swollen.  Mesosternum  and  metasternum  strongly  sulcate  throughout. 
Dorsal  ridge  of  metathoracic  peritreme  bilobed,  anterior  lobe  much  the  larger.  Corium  with  apical  margin 
straight.  Metathoracic  wing  with  antevannal  vein  present.  Anterior  and  intermediate  femora  with 
subapical  spines  beneath,  posterior  femur  with  three  major  subapical  spines  and  some  minor  ones  and  with 
basal  tubercle  well  developed.  Posterior  coxae  separated  by  about  half  the  width  of  a  coxa. 

Abdominal  sterna  III- VII  produced  into  broad,  triangular  spines,  lateral  margins  almost  smooth. 
Spermatheca  (Fig.  72)  with  bulb  lunate,  duct  long  and  tightly  convoluted.  Sclerites  of  wall  of  gynatrium 
triradiate,  with  three  arms  of  comparable  length,  one  arm  rising  to  meet  upper  ends  of  rami,  one 
descending  almost  to  meet  base  of  second  valvula  and  one  extending  horizontally  posteriorly.  Ovipositor 
with  valvulae  longer  and  more  slender  than  in  related  genera.  Male  genital  capsule  (Fig.  69)  short, 
posteriorly  emarginate,  emargination  filled  by  apices  of  the  short  parameres  (Figs  70, 71).  Aedeagus  (Figs 
66, 67, 68)  showing  phallotheca  of  form  typical  in  the  tribe;  conjunctiva  with  dorsomedian  lobe  M-shaped, 
membranous,  distal  dorsomedian  lobe  membranous,  small,  rounded-conical;  distal  dorsolateral  lobes 
small,  weakly  sclerotized,  apical  ventral  lobes  paired,  membranous,  subglobular,  distal  ventrolateral  lobes 
large,  membranous,  bilobed;  ventral  wall  of  conjunctiva  with  two  longitudinal  tracts  of  denticles  that 
extend  anterolaterally  up  sides  of  conjunctiva;  vesica  protected  at  base  by  a  dorsal,  cup-like  sclerite; 
ejaculatory  reservoir  with  long  wings  that  extend  inside  the  distal  dorsolateral  lobes  to  their  apices,  without 
straps. 

REMARKS.  The  anterolaterally  directed  pronotal  angles  are  shared  with  the  African  genus  My  la  and  the 
Oriental  Pseudomyla,  both  of  which  have  decumbent,  scale-like  pubescence  in  contrast  to  the  suberect  to 
erect  hairs  of  Neomevaniomorpha.  The  raised,  white  tip  of  the  scutellum  that  is  characteristic  of 
Psilolomia,  Mevaniomorpha  and  Mevanidea  is  not  found  in  this  genus;  it  is  least  well  developed  in 
Psilolomia  but  the  form  of  the  conjunctiva  is  dissimilar.  Probably  the  closest  relatives  of  the  genus 
Neomevaniomorpha  are  Mevaniomorpha  and  Mevanidea. 

DISTRIBUTION.  Central  Afro  tropical  region. 

Neomevaniomorpha  rodhaini  (Schouteden)  comb  n.,  stat.  n. 

(Figs  65-72) 

Mevaniomorpha  annulipes  rodhaini  Schouteden,  1938:  296.  LECTOTYPE  cf,  ZAIRE  (MRAC),  here 
designated  [examined] . 


184 


65 


W.  R.  DOLLING 


68 


72 


Figs  65-72  Neomevaniomorpha  rodhaini.  65,  dorsal  view  of  pronotum;  66,  dorsal  view  of  aedeagus;  67, 
lateral  view  of  phallotheca,  conjunctiva  and  vesica;  68,  ventral  view  of  conjunctiva  and  vesica;  69,  lateral 
view  of  genital  capsule;  70,  ventrolateral  view  of  left  paramere;  71,  dorsomedial  view  of  left  paramere; 
72,  spermatheca. 


Length:  cf ,  7-4-7-7 mm;  $,7-9-9-0 mm. 

Antennal  segment  I  about  equal  to  width  of  head  across  antennifers,  bearing  some  granules  about  as 
high  as  wide  or,  in  one  male  paralectotype,  with  a  few  tubercles  slightly  higher  than  wide  among  the 
granules.  Ratio  of  lengths  of  antennal  segments  about  1-00:0-96: 1-08:0-96.  Ratio  of  lengths  of  rostral 
segments  about  l-00:0-92:0-64:0-87. 

Shape  of  pronotum  as  in  Fig.  65.  Corium  with  apex  reaching  or  almost  reaching  suture  between 
later otergites  V  and  VI.  Sulcus  of  mesosternum  and  metasternum  very  broad.  Anterior  and  intermediate 
femora  each  with  a  single  subapical  spine  beneath;  posterior  femur  with  first  major  spine  less  than  half  as 
long  as  the  other  two,  followed  by  about  three  very  small  minor  spines,  space  between  the  two  larger  spines 
with  two  or  three  minor  spines  and  an  apical  series  of  four  minor  spines  present.  Length  of  posterior  tibia 
divided  by  length  of  posterior  femur  0-80-0-84. 

Abdominal  sternites  III- VII  with  their  posterolateral  angles  drawn  out  into  spines  equal  in  length  at 
most  to  one-third  of  the  width  of  their  respective  laterotergites.  Spermatheca  and  male  genitalia  as  in  Figs 
66-72  (see  description  of  genus  for  discussion  of  salient  features). 

Head  granulate;  gular  region  punctate;  vertex,  frons,  tylus  and  juga  shortly  tuberculate.  Pronotum 
granulate-punctate,  declivent  area  and  lateral  margins  tuberculate,  pleura  granulate-punctate,  sterna 
granulate,  scutellum  granulate-punctate  with  the  granules  on  its  lateral  margins  prominent.  Hemelytra 
with  clavus  and  corium  punctate  throughout,  veins  anteriorly,  especially  costal  margin  of  corium,  strongly 
granulate.  Legs,  especially  femora,  abdominal  sternites  and  laterotergites  granulate. 

Pubescence  of  moderate  to  rather  short,  suberect  hairs;  dorsum  of  head  and  declivent  part  of  pronotum 
with  long,  erect  hairs. 

Coloration  generally  mid-brown;  ventral  midline  of  thorax  and  of  abdomen  piceous.  Membrane  of 
hemelytra  fuscous,  the  veins  darker  with  occasional  small,  white  spots;  posterior  basal  angle  white, 
appearing  as  a  conspicuous,  white  triangle  just  behind  end  of  claval  commissure  when  hemelytra  are  at 
rest. 

REMARKS.  This  is  the  only  species  of  the  genus.  It  seems  strange  that  Schouteden  could  have  described  this 
insect  as  a  form  of  Mevaniomorpha  annulipes,  from  which  it  differs  strikingly  in  the  shape  of  the  pronotum 
and  in  coloration. 


TRIBE  PSEUDOPHLOEINI  185 

DISTRIBUTION.  Zaire  and  adjacent  part  of  Zambia. 

MATERIAL  EXAMINED 

Zaire:  1  cf  (lectotype),  Sankisia,  1911  (Dr  Rodham)  (MRAC). 

Zaire:  1  cf ,  Lulua,  Kapanga,  viii.1932  (F.  G.  Overlaet);  1  <f>,  Lulua,  Kapanga,  ix.1932  (F.  G.  Overlaet) 
(both  paralectotypes;  2  cf ,  Sankuru,  M'Pemba  Zeo  (Gandajika),  28.ix.1958  (R.  Marechat);  1  cf ,  Tshupa, 
Lukolela  (De  Guide);  1  $,  Lulua,  Kapanga,  i.1933  (F.  Overlaet);  1  <j>,  Wombali,  viii.1913  (P.  Vanderijst) 
(labelled  'allotypus')  (MRAC).  Zambia:  1  $,  Upper  Kalungwisi  Valley,  4200 ft  [1260 m],  11. ix.  1908  (5.  A. 
Neave)  (UM). 

MEVANIOMORPHA  Reuter 

Mevaniomorpha  Reuter,  1883: 13.  Type-species:  Mevaniomorpha  annulipes  Reuter,  by  monotypy. 

Body  oblong,  slightly  depressed;  connexivum  broad. 

Head  about  as  long  as  pronotum,  granulate-tuberculate.  Antennifers  strongly  divergent,  outer  apical 
process  porrect.  Antennal  segment  I  weakly  clavate,  granulate  or  granulate-tuberculate,  longer  or  shorter 
than  head;  III  longer  than  II,  both  slender,  granulate,  IV  shortest,  narrowly  fusiform,  specialized  sensory 
area  occupying  four-fifths  to  five-sixths  of  its  length.  Bucculae  occupying  about  one-quarter  of  ventral 
midline  of  head.  Rostrum  at  rest  reaching  to  disc  of  metasternum,  its  first  two  segments  subequal  in  length, 
fourth  shorter,  third  shortest  of  all. 

Pronotum  rather  shallowly  declivent,  granulate-tuberculate  and  punctate,  its  posterolateral  angles 
moderately  produced  laterally  with  posterolateral  spines  arising  abruptly  from  them  or  greatly  produced 
laterally  and  tapering  gradually  into  the  laterally-directed  spines;  lateral  margins  with  several  outstanding 
tubercles;  posterior  margin  straight,  smooth  in  front  of  scutellum,  granulate  further  towards  sides  of  body, 
prescutellar  spines  absent  but  granules  present  in  their  usual  position  on  each  side.  Scutellum  equilateral, 
its  apex  elevated  into  a  small,  white  swelling  occupying  about  one-fifth  of  its  length.  Mesosternum  and 
metasternum  longitudinally  sulcate  throughout.  Metathoracic  scent-gland  peritreme  with  dorsal  ridge 
bilobed,  anterior  lobe  much  the  larger,  constriction  between  lobes  shallow,  sometimes  obsolete.  Corium 
with  costal  and  apical  margins  slightly  convex.  Metathoracic  wing  with  antevannal  vein.  Anterior  and 
intermediate  femora  with  one  or  more  small  subapical  spines  beneath;  posterior  femur  with  three  major 
subapical  spines,  some  minor  ones  and  a  basal  tubercle.  Posterior  tibia  slightly  more  than  four-fifths  as  long 
as  femur.  Posterior  coxae  separated  by  rather  less  than  width  of  one  coxa. 

Abdominal  sternites  III-VII  with  posterolateral  angles  projecting  as  broad,  triangular  teeth,  lateral 
margins  almost  smooth.  Spermatheca  with  bulb  lunate,  duct  tightly  convoluted  within  concavity  of  bulb. 
Sclerites  of  wall  of  gynatrium  L-shaped.  Male  genital  capsule  posteriorly  emarginate,  emargination  filled 
by  apices  of  parameres.  Phallotheca  with  ventral  sclerite  short  ventrally,  strongly  produced  posterol- 
aterally,  dorsal  sclerites  very  weakly  sclerotized.  Conjunctiva  with  flat-topped  dorsomedian  lobe,  conical 
distal  dorsomedian  lobe,  short  distal  dorsolateral  lobes,  broad  apical  ventral  lobes,  large  distal  dor- 
soventral  lobes,  all  of  them  membranous;  two  strips  of  denticles  present  in  ventral  wall  of  conjunctiva; 
coiled  base  of  vesica  protected  by  two  sclerites  of  which  the  left  sclerite  is  twice  as  long  as  the  right  one. 

REMARKS.  This  genus  differs  from  Psilolomia  in  the  greater  projection  of  the  posterolateral  angles  of  the 
abdominal  sternites  and  the  presence  of  sclerites  at  the  base  of  the  vesica.  The  head  and  first  antennal 
segment  are  not  nearly  so  spiny  as  in  Mevanidea. 

DISTRIBUTION.  Tropical  Africa. 

Key  to  species 

1  Antennal  segment  I  shorter  than  width  of  head  including  eyes;  posterolateral  spines  of 
pronotum  arising  abruptly  from  the  posterolateral  angles;  width  across  apices  of  spines  less 
than  2-5  times  width  of  head  including  eyes.  (Widespread  in  tropical  Africa) annulipes(p.  185) 

-  Antennal  segment  I  longer  than  width  of  head  including  eyes ;  posterolateral  angles  of  pronotum 
tapering  gradually  into  spines;  width  across  apices  of  spines  more  than  2-5  times  width  of  head 
including  eyes.  (Forestsof  Central  and  West  Africa) picta(p.  187) 

Mevaniomorpha  annulipes  Reuter 

(Figs  73, 75,  76) 

[Clavigralla  elevator  (Fabricius)  sensu  Dallas,  1852:  511.  Misidentification;  not  Coreus  elevator  Fabricius, 
1803: 194.] 


186  W.  R.  DOLLING 

Mevaniomorpha  annulipes  Reuter,  1883:  13-14.  Holotype  $,  GHANA  (ZMU)  [examined]. 
[Mevaniomorpha  annulipes  subsp.  picta  Schouteden  sensu  Linnavuori,  1970:  43-45.  Misidentification.] 

Length:  cT,  7-2-8-2  mm;  $,  7-8-9-9  mm. 

Antennal  segment  I  about  as  long  as  width  of  head  across  antennifers.  Ratio  of  lengths  of  antennal 
segments  in  male  about  l-00:l-04:l-08:0-88,  in  female  about  l-00:l-04:l-12:0-84;  total  length  of  antenna 
less  than  3-7  times  width  of  head  including  eyes.  Ratio  of  lengths  of  rostral  segments  about 
l-00:0-92:0-54:0-78. 

Posterolateral  angles  of  pronotum  (Fig.  73)  prominent,  posterolateral  spines  arising  abruptly  from 
them,  directed  laterally;  width  across  apices  of  spines  divided  by  width  of  head  including  eyes  2-2-2-4. 

Aedeagus  as  in  Figs  75,  76  (see  description  under  genus). 

Head  dorsally  granulate-tuberculate,  laterally  granulate,  ventrally  punctate.  Pronotum  granulate 
throughout,  tuberculate  anteriorly,  punctate  posteriorly;  scutellum  and  pleura  punctate-granulate;  thor- 
acic and  abdominal  sterna  and  laterotergites  weakly  granulate.  Clavus  and  corium  punctate  throughout, 
veins  sparsely  and  weakly  granulate;  femora  granulate. 

Pubescence  of  rather  short,  suberect  hairs,  almost  decumbent  on  head  and  pleura;  longer,  erect  hairs 
present  on  dorsum  of  head  and  pronotum. 

Colour  pattern  of  medium  brown  and  piceous  markings  on  a  paler,  yellow  ground.  The  most  prominent 
dark  markings  are  a  stripe  along  anterior  midline  of  pronotum,  a  pair  of  spines  in  front  of  anterolateral 
angles  of  pronotum,  apical  margin  of  corium,  spot  in  apical  part  of  disc  of  corium  usually  separate  from 
dark  apical  margin,  posterior  two-thirds  of  laterotergites  III-V,  posterior  margins  of  laterotergites  V  and 
VI,  anterior  margins  of  laterotergites  III- VII  and  extensive  mottling  on  apices  of  posterior  femora. 

REMARKS.  Some  specimens  in  the  north-eastern  part  of  the  range  have  shorter  posterolateral  pronotal  and 
abdominal  spines  than  the  main  population;  such  specimens  were  referred  to  the  variety  picta  Schouteden 
by  Linnavuori  (1970:  43-^45);  picta  in  fact  has  longer  spines  than  true  annulipes  and  appears  to  be  a  good 
species. 

DISTRIBUTION.  Widespread  in  tropical  Africa. 

MATERIAL  EXAMINED 

Ghana:  1  cf  (holotype),  Addah  (Reitter)  (ZMU). 

Guinea:  1  cf,  Nimba,  Yalanzou,  'ii.vi.42'  (Lamotte),  1  cf,  Nimba,  Keoulenta,  'ii.vi.42'  (Lamotte) 
(MNHN).  Sierra  Leone:  1  $  (leg.  Morgan  fide  Dallas,  1852:  511)  (labelled  '425a'  and  'Clavigralla 
elevator,');  1  cf ,  Mopeille,  l.vii.1926  (Hargreaves)  (BMNH).  Ivory  Coast:  1  $ ,  Tai,  12.xi.1979  (Couturier) 
(MNHN);  1  Cf ,  Bingerville,  xii.1963  (Decelle);  1  $,  Korea,  au  sud  de  Daloa,  ix.1961  (Decelle)  (MRAC). 
Nigeria:  1  cf,  Calabar,  9.xi.l955  (Bechyne,  Exped.  Mus.  G.  Frey);  1  cf,  Akpasha  Udi,  27.X.1955 
(Bechyne,  Exped.  Mus.  G.  Frey);  1  cf ,  near  Bida,  between  Dabba  and  Kutiwenji,  20. xi.  1970  (Deeming);  1 
$ ,  Gombe,  Matzoro  Lakes,  i.1929  (Lloyd)  (BMNH).  Cameroun:  2  cf ,  Batouri  District,  3°45'N,  13°45'E, 
750  m,  I.v-6.vi.l935  (Merfield)  (BMNH);  1  $,  Baigom,  reg.  Bamoun  (MNHN).  Zaire:  1  cf ,  Gamangui, 
ii.1910  (Lang  &  Chapin)  (AMNH);  1  cf,  Haut-Uele,  Manda,  18.iii.1925  (Schouteden);  1  cf,  Sankuru, 
M'Pemba  Zeo  (Gandajika),  1960  (Marechal);  1  cf,  Kivu,  Mulungu,  1939  (Hendrickx);  1  cf,  Kivu, 
Mulungu-Tshibinda,  xi.1951  (Lefevre);  1  cf,  Kivu,  Mwenga,  Kitutu,  rive  de  1'Elila,  650  m,  foret 
marecageuse  primaire,  humus,  iv.1958  (Leleup);  1  cf,  Katanga,  Luembe,  viii-ix.1956  (de  Caters);  1  $, 
Lulua,  Kafakumba,  xii.1932  (Overlaet);  1  $,  Maulema,  Wamaza,  iii-iv.1957  (Cotonco);  1  $,  Gandajika, 
iv.1959  (Decelle);  1  $,  Bas-Congo,  Thysville,  1959-1963  (Michaux);  1  $,  Bas-Congo,  Mayidi,  1942  (van 
Eyen);  1  $,  Kivu,  Mulungu,  1938  (Hendrickx);  1  $,  Kivu,  Mulungu,  Tshibinda,  xi.1951  (Lefevre);  1  $, 
Kivu,  Ibanda,  1952  (Vandelannoite)  (MRAC).  Uganda:  1  cf,  Mutunda,  l.vi.1911  (Marshall);  1  $, 
Western  Ankole,  4500-5000  ft  (1350-1500  m),  10-14.X.1911  (Neave)  (BMNH).  Kenya:  1  cf,  Embu, 
12.ix.1914  (Browne);  1  $,  N.  slopes  of  Mt  Kenya,  on  Embu-Meru  road,  4500-5000  ft  (1350-1500  m), 
13-14.ii.1911  (Neave);  1  $,  Ngong,  v.1926  (Gedye);  1  cf ,  1  $,  Kaimosi,  iii-iv.1932  (Turner);  1  £,  Chyulu 
Hills,  5200  ft  (1600  m),  iv.1938  (Coryndon  Mus.  Exped.)  (BMNH).  Zambia:  1  cf ,  Lake  Bangweulu, 
N'Sumba  Island,  25.xi.1946,  beaten  from  climbers  (Steele);  1  £,  Lake  Bangweulu,  Kapola,  N.  of  Kapata, 
27.X.1946  (Steele);  2  cf ,  1  $,  Lake  Bangweulu  district,  N'Salushi  Island,  13.xi.1946  (Steele);  4  cf,  Upper 
Luanga  River,  27.vii-13.viii.  1910  (Neave);  1  9,  Luangwa  to  Petauke,  14-17.ix.1910  (Neave)  (BMNH);  1 
Cf ,  Lower  Kalungwisi  valley,  3500  ft  (1050  m),  dense  forest,  12-13. ix.1908  (Neave)  (UM).  Malawi:  1  $, 
Ruo  valley,  1000-2000  ft  (300-600  m),  21-25.iv.1910  (Neave)  (BMNH).  South  Africa:  1  $,  Transvaal, 
Louis  Trichard,  20-30. xii.  1956  (Capener)  (J.  A.  Slater  coll.);  1  $  Natal,  'P.  town'  (?=Pinetown),  Gramin 
(BMNH). 


TRIBE  PSEUDOPHLOEINI  187 

Mevaniomorpha  picta  Schouteden  stat.  n. 
(Fig.  74) 

Mevaniomorpha  annulipes  picta  Schouteden,  1938:  296.  LECTOTYPE  cf,  ZAIRE  (MRAC),  here  desig- 
nated [examined]. 

Length:  cf,  8-1-8-4  mm;  $,8-6-9-3  mm. 

Antennal  segment  I  longer  than  width  of  head  including  eyes;  ratio  of  lengths  of  antennal  segments 
about  1 -00:0-91 :0-96:0-71;  total  length  of  antennae  more  than  4-1  times  width  of  head  including  eyes.  Ratio 
of  lengths  of  rostral  segments  about  l-00:0-93:0-55:0-79. 

Pronotum  (Fig.  74)  with  posterolateral  angles  merging  gradually  into  posterolateral  spines,  width  across 
apices  of  spines  2-6-3-0  times  width  of  head  including  eyes;  tubercles  of  disc  and  margins  long. 

Male  genitalia  similar  to  those  of  M.  annulipes  but  ejaculatory  reservoir  complex  with  wings  rather 
longer. 

Pubescence  as  in  M.  annulipes;  granulation  and  tuberculation  more  pronounced.  Colour  pattern  very 
similar  to  that  of  annulipes  but  all  elements  of  pattern  darker;  antennae  rather  dark  brown  throughout. 

REMARKS.  This  species  differs  from  M.  annulipes  in  the  longer  appendages,  more  pronounced  posterolat- 
eral spines  of  the  pronotum  and  darker  colour.  Linnavuori  (1970:  43-45),  presumably  basing  his 
identification  on  coloration,  referred  dark  specimens  of  true  annulipes  to  this  taxon;  these  specimens  in  fact 
had  slightly  shorter  spines  and  appendages  than  typical  annulipes.  After  the  above  description  was 
completed,  Dr  V.  van  Zeijst  showed  me  two  males  of  picta  from  the  Tai  forest  in  the  Ivory  Coast;  these 
specimens  had  very  long,  slender  pronotal  spines  and  were  almost  black;  superficially,  they  closely 
resembled  the  unrelated  Clavigralla  hystrix  Dallas,  which  occurred  in  the  same  forest. 

DISTRIBUTION.  Rain  forests  of  Central  and  West  Africa. 

MATERIAL  EXAMINED 

Zaire:  1  cf  (lectotype),  Sankuru,  Komi,  iv.1930  (/.  Ghesquiere)  (MRAC). 

Zaire:  1  cf,  Sankuru,  Komi,  2.V.1930  (Ghesquiere);  1  cf,  Stanleyville  (=  Kisangani),  20.xii.1929 
(Collart);  1  $,  Uele,  Dingila,  ix.1933  (Bredo);  1  $,  Lomani,  Kambaye,  vii.1930  (Quarre)  (paralectotypes) 
(MRAC);  1  $,  Yangambi,  xi.1937  (Henrard)  (MRAC).  Cameroon:  1  $,  Batouri  (MNHN). 

MEVANIDEA  Reuter 

Mevania  Stal,  1866:  110.  Type-species:  Clavigralla  spiniceps  Signoret,  by  monotypy.  [Homonym  of 

Mevania  Walker,  1854:  442-^43.] 

Mevanidea  Reuter,  1883: 11-12.  Type-species:  Mevanidea  granulifera  Reuter,  by  monotypy. 
Mevaniella  Bergroth,  1907:  146.  [Replacement  name  for  Mevania  Stal.]  Syn.  n. 

Body  not  depressed,  conexivum  moderately  expanded.  Aspect,  especially  of  head  and  pronotum,  spinose 
(Fig.  77). 

Head  slightly  shorter  than  pronotum,  dorsally  with  numerous,  long,  spine-like  tubercles.  Eyes  small, 
prominent.  Antennifers  moderately  divergent,  external  apical  processes  porrect  and  directed  slightly 
downwards.  Antennal  segment  I  strongly  clavate,  bearing  numerous  spine-like  tubercles  on  its  expanded 
apical  part.  Antennal  segments  I  and  IV  subequal  in  length,  IV  fusiform;  II  subequal  to  or  rather  longer 
than  I,  III  much  the  longest;  II-IV  almost  smooth,  bearing  hairs  about  as  long  as  width  of  segments  II  and 
III.  Bucculae  occupying  about  one-quarter  of  ventral  midline  of  head.  Rostrum  at  rest  reaching  to  anterior 
margin  or  disc  of  metasternum;  segment  II  slightly  and  IV  distinctly  shorter  than  I,  III  obviously  shortest. 

Pronotum  strongly  declivent,  posterior  margin  straight  or  slightly  convex,  prescutellar  spines  well 
developed,  tubercles  present  between  them  and  posterolateral  angles,  along  lateral  margins  and  on  disc, 
including  some  on  posterior  part.  Posterolateral  angles  of  pronotum  slightly  produced,  posterolateral 
spines  arising  abruptly  from  them,  directed  laterally  and  slightly  anteriorly.  Scutellum  equilateral,  slightly 
convex,  its  apical  one-third  conspicuously  swollen,  china- white.  Mesosternum  deeply  sulcate  throughout; 
metasternum  sulcate  for  half  or  all  of  its  length.  Metathoracic  peritreme  with  its  dorsal  ridge  simple  or 
unequally  bilobed,  the  anterior  lobe  in  the  latter  case  much  the  larger.  Metathoracic  wing  with  antevannal 
vein.  Anterior  and  intermediate  femora  with  two  rows  of  tubercles  and  granules  beneath.  Posterior  femur 
with  basal  tubercle  and  beneath  with  three  major  subapical  spines,  the  first  spine  two-thirds  as  long  as  the 
others;  usually  three  minor  spines  between  the  two  biggest  ones  and  an  apical  series  of  four.  Posterior  tibia 
short,  two-thirds  to  three-quarters  as  long  as  femur  and  conspicuously  curved  near  base. 

Abdominal  sternites  III-VI  with  posterolateral  angles  produced  into  broad,  triangular  spines,  lateral 


188 


W.  R.  DOLLING 


73 


Figs  73-78  73,  Mevaniomorpha  annulipes,  dorsal  view  of  head,  pronotum,  scutellum  and  first  antennal 
segment.  74,  M.  picta,  dorsal  view  of  head,  pronotum  and  first  antennal  segment.  75, 76,  M.  annulipes: 
(75)  ventral  view  of  conjunctiva  and  vesica;  (76)  dorsal  view  of  phallotheca,  conjunctiva  and  vesica.  77, 
Mevanidea  hystrix,  dorsal  view  omitting  legs  and  last  three  antennal  segments.  78,  Arenocoris  interme- 
dius,  antenna. 


margins  of  sterna  granulate.  Spermatheca  with  bulb  very  narrow,  duct  three  to  four  times  as  long  as  bulb, 
convoluted.  Male  conjunctiva  with  dorsomedian  lobe  M-shaped,  membranous,  distal  dorsomedian  lobe  of 
similar  size,  also  membranous,  distal  dorsolateral  lobes  obsolete,  distal  ventrolateral  lobes  large,  bifid, 
weakly  sclerotized  on  their  posterior  faces,  apical  ventral  lobes  membranous,  paired,  ventral  wall  of 
conjunctiva  with  or  without  a  pair  of  toothed,  sclerotized  strips;  basal,  coiled  part  of  vesica  protected  by 
two  subequal  sclerites. 


TRIBE  PSEUDOPHLOEINI  189 

REMARKS.  The  appearance  of  the  two  species  of  this  genus  is  very  characteristic  (Fig.  77),  the  spiny  head 
and  first  antennal  segment  being  particularly  notable.  The  genus  is  probably  derived  from  a  Mevaniomor- 
p/io-like  form. 

DISTRIBUTION.  Africa  and  Madagascar. 

Key  to  species 

1    Membrane  of  fore  wing  with  an  irregular,  opaque,  piceous  or  dark  brown  spot  occupying  about 

half  of  its  area.  (Africa) hystrix(p.  189) 

-    Membrane  of  forewing  translucent,  with  some  darker  markings  along  veins  and  sometimes  with 
dark  amber  shading  of  area  between  veins,  but  never  with  an  opaque  spot.  (Madagascar) 

spiniceps(p.  190) 

Mevanidea  hystrix  (Gerstaecker) 
(Fig.  77) 

Mevania  hystrix  Gerstaecker,  1873:  408-409.  Holotype  cf ,  KENYA  (MNHU)  [examined]. 

Mevanidea  granulifera  Reuter,  1883:  12.  Holotype  $,  GHANA  (ZMU)  [examined].  [Synonymized  by 

Linnavuori,  1978:  36?] 

Mevanidea  kilimana  Schouteden,  1910: 157.  [Nomen  nudum.] 
Mevaniella  kilimana  Schouteden,  1912:  55.  LECTOTYPE  cf ,  KENYA  (NR),  here  designated  [examined]. 

[Synonymized  by  Schouteden,  1938:  295.] 
Mevanidea  hystrix  (Gerstaecker)  Schouteden,  1938:  295. 

Length:  cf ,  5-7-6-8 mm;  $,6-2-7-6 mm. 

Ratio  of  lengths  of  antennal  segments  in  male  about  1-00: 1-02: 1-30: 1-02,  in  female  about 
1-00:1-14:1-41:1-00.  Ratio  of  lengths  of  rostral  segments  in  male  about  l-00:0-97:0-55:0-79,  in  female 
about  l-00:0-97:0-50:0-75.  Head  dorsally  with  numerous  granules  and  tubercles  of  various  lengths,  most 
noticeably  a  pair  of  spine-like  tubercles  close  together  between  eyes  with  a  third  immediately  behind  them, 
a  long,  spine-like  tubercle  on  each  side  of  vertex  just  posterior  to  inner  basal  angles  of  antennifers,  a 
staggered  row  of  long  tubercles  all  along  tylus  and  others,  rather  shorter,  on  juga;  sides  and  ventral  surface 
of  head  punctate-granulate. 

Pronotum  with  lateral  margins  almost  straight,  bearing  three  or  four  long  tubercles,  lateral  angles 
suddenly  prominent;  prescutellar  spines  long,  slender,  sinuous;  centre  of  disc  with  a  widely  spaced  pair  of 
conical  tubercles,  two  groups  of  partially  fused,  blunt  granules  behind  them  but  less  widely  spaced  and  a 
row  of  three  similar  clusters  of  granules  between  these  and  posterior  margin  in  addition  to  more  generally 
distributed  granules  and  tubercles.  Scutellum  and  thoracic  pleura  and  sterna  punctate-granulate.  Hemely- 
tra  with  clavus  and  corium,  except  for  smooth  area  in  middle  of  its  apical  part,  punctate,  veins  anteriorly 
and  basally  granulate.  Femora  strongly  granulate,  shortly  tuberculate  dorsally  and  ventrally,  with  one  or 
two  subapical  tubercles  of  the  anterior  and  intermediate  femora  enlarged  and  occupying  the  usual  positions 
of  subapical  spines;  posterior  femur  with  three  major  subapical  spines  beneath,  one  to  four  minor  spines  in 
the  space  between  the  first  two  and  three  or  four  in  the  space  between  the  middle  and  last,  with  an  apical 
series  of  three  or  four.  Posterior  tibia  about  two-thirds  as  long  as  femur. 

Ventral  wall  of  conjunctiva  of  aedeagus  with  a  pair  of  apically  divergent,  toothed,  sclerotized  strips; 
vesica  short,  about  as  long  as  either  of  the  wings  of  the  ejaculatory  reservoir  complex. 

Pubescence  throughout  of  moderate  to  short,  suberect  hairs  with  some  long,  erect  hairs  on  pronotum. 

Colour  pattern  as  in  Fig.  77;  the  pale  areas  being  stramineous  and  the  dark  areas  varying  from  dark 
brown  to  dark  red  in  different  specimens.  The  most  striking  features  of  the  colour  pattern  are  the 
china- white,  swollen  apex  of  the  scutellum,  an  irregular,  opaque,  dark  brown  blotch  in  the  middle  of  the 
membrane  of  the  hemelytra  and  a  dark  brown  (never  red)  transverse  spot  near  middle  of  corium  edged 
proximally  with  white;  laterotergites  III  to  VI  largely  dark  with  a  pale  band  anteriorly,  VII  largely  pale. 

REMARKS.  I  can  trace  no  earlier  synonymy  of  granulifera  with  hystrix  than  that  of  Linnavuori  (1978:  36); 
however,  the  synonymy  is  not  indicated  as  new  in  that  publication.  The  holotype  of  M.  hystrix  Gerstaecker 
is  a  small  (6-0  mm)  and  very  red  male  from  Mombasa  (Kenya)  and  that  of  M.  granulifera  Reuter  a  large  (7-1 
mm)  blackish  brown  female  from  'Addah'  (i.e.  Ada,  Ghana).  The  great  distance  (4500  km)  between  the 
type-localities  and  the  striking  difference  between  specimens  at  the  extremes  of  the  range  of  colour 
variation  have  led  earlier  workers  to  recognize  two  species  on  the  African  continent.  The  reddish 
coloration  is  in  general  more  pronounced  in  specimens  from  eastern  and  southern  Africa  and  the  brownish 


190  W.  R.  DOLLING 

coloration  more  pronounced  in  those  from  western  Africa.  The  dark  brown  blotch  in  the  middle  of  the 
membrane  of  the  hemelytra  is  variable  in  extent  but  not  in  colour.  The  egg  of  this  species  is  illustrated  by 
Cobben  (1968:  97,  fig.  84). 

DISTRIBUTION.  Widespread  in  subsaharan  Africa. 

MATERIAL  EXAMINED 

Kenya:  1  cf  (holotype  of  hystrix),  Mombasa  (van  der  Deckeri)  (MNHU).  Ghana:  1  9  (holotype  of 
granulifera),  Addah  (Reitter)  (ZMU).  Kenya:  1  $  (lectotype  ofkilimana),  Kilimanjaro  (Sjoestedt)  (NR). 

Sierra  Leone:  1  $,  Port  Lokko,  30.viii.1925  (Hargreaves)  (BMNH).  Ivory  Coast:  1  $,  Adiopodoume, 
iv-v.1964  (Cobben)  (BMNH);  1  $,  Bouake,  ii.1963  (Schmitz)  (MRAC).  Ghana:  1  cf,  Kintampo, 
7.xii.l965  (Leston)  (BMNH).  Nigeria:  1  cf,  Kumba,  15.xi.1955  (Bechyne)  (BMNH).  Sudan:  1  cf, 
Ingessana  Mts,  Blue  Nile,  18-22.xi.1982  (Linnavuori)  (BMNH).  Zaire:  1  cf ,  Yangambi,  xii.1959  (De- 
celle);  1  $,  Muranan  'Deke,  v.1953  (De  Francqueri)  (MRAC);  1  cf,  Mpese,  ll.vi.1937  (Cooremari) 
(IRSNB);  7  Cf,  5  $,  Faradje,  29°40'E,  3°40'N,  i.1913  (Lang  &  Chapiri)  (AMNH).  Uganda:  1  cf ,  Bussu 
Busoga,  v.1909  (Bayon);  1  $,  Chugwe,  Mabira  Forest,  3500-3800  ft  (ca  1100  m),  16-25.vii.1911  (Neave) 
(BMNH).  Kenya:  1  cf,  Rabai,  vii.1937  (van  Someren);  1  cf,  Mombasa,  iii.1921  (van  Someren);  1  <j>, 
Chyula  Hills,  5200  ft  (1600  m),  iv.1938  (Coryndon  Museum  Expedition)  (BMNH).  Tanzania:  1  cf ,  Uvira, 
28-29.viii.1931  (Ogilvie);  1  cf,  Zanzibar,  near  Mazi  Moja,  20.viii-ll.ix.1924  (Snelt)  (BMNH);  1  $, 
Zanzibar  (Ashby)  (AMNH).  Malawi:  1  cf ,  between  Fort  Mangoche  and  Chikala  Boma,  about  4000  ft 
(1200  m),  20-25.iv.1910  (Neave);  1  $,  Mlanje,  2.xii.l912  (Neave)  (BMNH).  Zambia:  2  cf,  2  $,  Upper 
Luanga  River,  27.vii-13.viii.1910  (Neave);  1  $,  Lake  Bangweulu  district,  ix-xii.1946  (Steele);  1  $,  Lake 
Bangweulu,  Chiluwi  Island,  beaten  from  lemon  (Citrus  limon),  29. xi. 1946  (Steele)  (BMNH).  South  Africa: 
1  Cf ,  1  $ ,  Transvaal,  Kruger  National  Park,  3  miles  E.  of  Satara  Camp,  Nwandzi  River,  29. iv. 1968  (Schuh, 
Slater  &  Sweet)  (J.  A.  Slater  collection);  1  cf ,  Transvaal,  Johannesburg,  6000  ft  (1800  m),  ii.1895  (Cregoe); 
1  $,  Transvaal,  Johannesburg,  6000  ft  (1800  m),  iv.1899  (Cregoe);  1  cf,  Natal,  Estcourt,  i.1897;  2  cf, 
Natal,  Tugela  River,  near  Weenen,  14.iii.1897;  1  $,  Natal,  Durban,  1874  (Bell-Marley)  (BMNH). 

Mevanidea  spiniceps  (Signoret)  comb.  n. 

Clavigralla  spiniceps  Signoret,  1861:  944.  LECTOTYPE  $,  MADAGASCAR  (NMV),  here  designated 

[examined]. 

Mevania  spiniceps  (Signoret)  Stal,  1866: 110-111. 
Mevaniella  spiniceps  (Signoret)  Bergroth,  1907: 146. 

Length:  cf,  6-8-7-8  mm;  $,7-3-8-2  mm. 

Ratio  of  lengths  of  antennal  segments  in  male  about  1-00:1-30:1-55:1-15,  in  female  about 
1-00:1-14:1-48:1-04.  Ratio  of  lengths  of  rostral  segments  in  male  about  l-00:0-92:0-54:0-87,  in  female 
about  l-00:0-92:0-55:0-78.  Head  strongly  spiny-tuberculate  but  with  fewer  tubercles  and  granules  than  M. 
hysterix;  the  pair  of  long  tubercles  on  vertex  between  eyes  without  a  tubercle  behind  them.  Posterior  tibia 
about  three-quarters  as  long  as  femur. 

Aedeagus  with  no  ventral  sclerotized,  toothed  strips  on  conjunctiva  and  with  vesica  about  twice  as  long 
as  either  wing  of  ejaculatory  reservoir. 

Colour  pattern  of  dark  to  pale,  often  somewhat  rufous  (but  never  strongly  reddish)  brown  areas  and 
stramineous  areas;  membrane  of  hemelytra  without  opaque,  brown  blotch  though  with  some  brown 
markings  along  veins  and  occasionally  with  some  dark  amber  shading  of  membrane  between  veins. 
Laterotergites  III  to  V  largely  dark,  each  with  a  pale  band  anteriorly,  VI  and  VII  largely  pale. 

REMARKS.  This  species  resembles  M.  hystrix  in  general  appearance.  The  main  differences  are  in  the 
aedeagus,  where  the  longer  vesica  of  spiniceps  is  presumably  plesiomorphic  and  the  absence  of  ventral 
sclerotized  strips  in  the  conjunctival  wall  apomorphic,  and  the  absence  of  the  dark  brown  blotch  on  the 
hemelytral  membrane.  The  presence  of  only  two  spines,  rather  than  three,  on  the  vertex  between  the  eyes 
is  constant  in  all  the  specimens  examined.  M.  spiniceps  and  a  few  species  of  Clavigralla  (tribe  Clavigrallini) 
are  the  only  Pseudophloeinae  known  from  Madagascar. 

DISTRIBUTION.  Madagascar. 

MATERIAL  EXAMINED 

Madagascar:  1  $  (lectotype)  (NMV). 

Madagascar:  1  $,  Tulear  Province,  Tongobory,  200  m,  27.iii.1968  (K.M.G.  &  P.D.)  (BMNH);  4  cf ,  2 
$ ,  [no  precise  locality],  1930  (Sicard);  2  cf ,  Baie  de  Baly,  Plateau  de  Soalala,  1930  (Joly);  2  cf ,  no  data;  1 
Cf,  Region  de  Sud-est,  Fort-Dauphin,  i.1901  (Alluaud);  1  cf,  Nossi-Be,  5.vii.l900  (Alluaud);  1  cf, 


TRIBE  PSEUDOPHLOEINI  191 

Ivondro,  vii.1900  (Alluaud);  1  cf  [no  precise  locality],  (Fairmaire);  1  $,  Region  de  Sud-est,  Vallee  du 
Fanjahira,  xii.1901  (Alluaud);  1  $,  Vallee  du  Fanjahira,  Isaka,  xii.1901  (Alluaud);  1  $,  Nossi-Be,  Foret 
Loukoube,  1897  (Alluaud);  1  $,  Sahambava,  Fianarantsoa  (MNHN). 

AtfEJVOCORISHahn 

Arenocoris  Hahn,  1834:  109-110.  Type-species:  Coreus  falleni  Schilling,  by  subsequent  designation  of 

Blanchard,  1842:  312. 

[Atractus  Laporte  sensu  Curtis,  1834:  legend  to  pi.  500.  Misidentification.] 
Pseudophloeus  Burmeister,  1835:  301,  302,  308.  Type-species:  Coreus  falleni  Schilling,  by  subsequent 

designation  of  Brulle,  1835:  359. 

Ammocoris  Agassiz,  1848:  48,  94.  [Unjustified  emendation  of  Arenocoris  Hahn.] 
Psammocoris  Marshall,  1868:  281.  [Unjustified  emendation  of  Arenocoris  Hahn.] 
Boudicca  Kirkaldy,  1909:  30.  [Unnecessary  replacement  name  for  Pseudophloeus  Burmeister.] 

Body  strongly  depressed,  ovate;  connexivum  considerably  expanded  in  middle.  Body  and  appendages 
conspicuously  granulate-tuberculate . 

Head  distinctly  longer  than  pronotum.  Antennifers  divergent  with  outer  apical  process  either  porrect  or 
deflexed.  Antennae  with  segment  I  much  shorter  than  width  of  head,  strongly  incrassate  except  for  basal 
one-eighth;  segments  II  and  III  slender,  II  much  shorter  than  I,  III  about  4  times  as  long  as  II,  IV  short, 
fusiform,  intermediate  in  thickness  between  I  and  II.  Bucculae  occupying  about  two-fifths  of  ventral  length 
of  head.  Rostrum  at  rest  reaching  to  posterior  margin  of  mesosternum. 

Pronotum  weakly  declivent,  abruptly  widened  posteriorly;  posterolateral  angles  truncate,  sometimes 
auriculate,  not  pointed  and  not  bearing  posterolateral  spines;  posterior  margin  almost  straight,  weakly 
developed  prescutellar  lobes  (not  spines)  present.  Scutellum  equilateral,  lateral  margins  elevated  anteri- 
orly, apex  elevated  and  bilobed.  Mesosternum  throughout  and  metasternum  anteriorly  deeply  sulcate  in 
midline.  Metathoracic  scent-gland  peritreme  with  dorsal  lobe  entire,  its  dorsal  margin  evenly  rounded, 
descending  anteriorly  and  posteriorly  to  enclose  orifice  in  at  least  its  dorsal  half.  Membrane  of  hemelytron 
with  venation  reticulate.  Metathoracic  wing  with  antevannal  vein  well  developed.  Anterior  and  intermedi- 
ate femora  without  subapical  spines  beneath,  sometimes  with  a  very  slightly  enlarged  tubercle  in  this 
position;  posterior  femur  with  a  single,  large  tubercle  or  blunt  spine  subapically  beneath,  very  rarely  with 
an  additional,  slightly  enlarged  tubercle  distad  of  this;  base  of  posterior  femur  with  a  prominent  tubercle 
adjacent  to  trochanter. 

Abdominal  margin  evenly  rounded,  posterolateral  angles  of  sternites  obtuse  or  right-angled,  not 
prominent.  Male  genital  capsule  short  with  tongue  triangular,  lip  obtusely  angled  in  middle;  parameres 
broad,  flat,  their  apices  filling  posterior  emargination  of  capsule.  Phallotheca  comprising  two  slender, 
dorsal  sclerites  and  a  slightly  shorter,  broad,  ventral  sclerite.  Conjunctiva  with  dorsomedian  lobe  broad, 
membranous;  distal  dorsolateral  lobes  very  long,  their  posteroventral  sides  supported  by  sclerotized 
extensions  of  wings  of  ejaculatory  reservoir  apparatus;  vesica  not  protected  basally  by  sclerites;  apical 
ventral  lobes  small,  membranous;  distal  ventrolateral  lobes  large,  bilobed,  the  lower  lobe  sclerotized  on  its 
posterior  face;  ventral  wall  of  conjunctiva  with  two  slender,  parallel  sclerites;  ejaculatory  reservoir 
complex  with  wings  long,  straps  absent.  Ovipositor  with  second  valvulae  emarginate  at  apex.  Sclerites  of 
dorsal  wall  of  gynatrium  short,  each  comprising  a  narrow,  longitudinally  elongate,  vertical,  mesal  plate 
supporting  a  thin,  sclerotized  ring  and  a  narrow,  horizontal,  transverse,  anterior  plate  articulating  laterally 
with  second  valvifer  and  its  ramus.  Spermatheca  with  bulb  lunate  or  half-moon  shaped,  duct  about  1-5 
times  as  long  as  bulb,  slightly  undulate  but  not  convoluted. 

INCLUDED  SPECIES.  Stichel  (1962:  194-195)  lists  six  species:  egenus  (Horvath,  1917),  falleni  (Schilling, 
1829),  gestroi  (Bergevin,  1930),  intermedius  (Yakovlev,  1883),  latissimus  Seidenstiicker,  1960  and  waltli 
(Herrich-Schaeffer,  1834).  Chernova  (1979:  579)  established  the  synonymy  of  egenus  with  falleni,  and  the 
synonymy  of  gestroi  with  intermedius  is  established  below. 

REMARKS.  This  is  essentially  a  Palaearctic  genus,  with  a  single  eremic  species  intrusive  into  the  north- 
eastern part  of  the  Afrotropical  region.  Within  the  tropical  fauna  it  is  readily  distinguished  from  other 
genera  of  Pseudophloeinae  by  the  very  short  antennal  segments  I  and  II. 

DISTRIBUTION.  Europe,  North  Africa,  Canary  Islands,  western  Asia;  mostly  eremic  but  two  species  extend 
into  cooler  parts  of  Europe  and  another  into  Ethiopia  and  Chad. 


192  W.  R.  DOLLING 

Key  to  species 

1  Outer  apical  process  of  antennifer  deflexed 

-  Outer  apical  process  of  antennifer  porrect 3 

2  Antennal  segment  I  longer,  length  of  incrassate  part  0-46  or  more  times  width  of  head  including 

eyes.  (Spain,  Portugal,  western  Asia  and  northern  Africa  southwards  to  Chad  and 

Ethiopia) intermedius(p.  192) 

-  Antennal  segment  I  shorter,  length  of  incrassate  part  less  than  0-46  times  width  of  head  including 

eyes.  (Europe,  western  Asia,  North  Africa  from  Morocco  to  Libya) fallen! (Schilling) 

3  Head  shorter,  its  length  equal  to  its  width  including  eyes.  (Europe,  Canary  Islands,  coastal 

countries  of  North  Africa,  western  Asia,  Central  Asia) waM(Herrich-Schaeffer) 

-  Head  longer,  its  length  about  1-2  times  its  width  including  eyes.  (Turkey  and  adjacent  areas  of 

U.S.S.R.) /atfssimusSeidenstucker 

Arenocoris  intermedius  ( Yakovlev) 
(Fig.  78) 

Pseudophloeus  intermedius  Yakovlev,  1883:  101-103.  Holotype  (sex  unknown),  U.S.S.R.:  Krasnovodsk 

(Bekker)  (IZ)  [not  examined]. 
Pseudophloeus  angustus  Reuter,  1891:  139.  Holotype  $,  EGYPT  (ZMU)  [examined].  [Synonymized  by 

Kirichenko,  1952: 165.] 
Pseudophloeus  gestroi  Bergevin,  1930:  32.  Holotype  cf ,  LIBYA:  Giarabub,  1926-1927  (Confalonieri) 

(Museo  Civico  di  Storia  Naturale,  Geneva)  [not  examined].  Syn.  n. 
Arenocoris  intermedius  (Yakovlev)  Kirichenko,  1952: 165. 

Length:  cf ,  6-0-7-2 mm,  $,5-6-7-4 mm. 

Head  granulate-tuberculate.  Length  of  head  equal  to  its  width  inclusive  of  eyes;  antennifers  strongly 
divergent,  their  outer  apical  processes  deflexed.  Antennae  (Fig.  78)  with  segment  I  abruptly  incrassate 
from  a  slender  base,  incrassate  part  tuberculate;  segments  II  and  III  slender,  weakly  and  densely  granulate; 
IV  shortly  ovate-fusiform,  nearly  as  thick  as  I  and  with  apical  sensory  area  occupying  about  one-half  of 
its  length.  Length  of  antennal  segment  I  (excluding  slender  basal  part)  divided  by  width  of  head  including 
eyes  0-47-0-61.  Ratio  of  lengths  of  segments  (again  excluding  slender  base  of  I)  in  male  about 
l-00:0-58:2-34:0-80,  in  female  about  l-00:0-57:2-25:0-83.  Ratio  of  lengths  of  rostral  segments  about 
l-00:0-98:0-52:0-72. 

Pronotum  granulate-tuberculate,  lateral  margins  each  bearing  three  or  four  prominent  tubercles,  disc 
with  two  longitudinal  rows  of  semiglobular  granules,  most  of  these  in  contact  with  the  neighbouring  ones, 
not  standing  separately;  anterolateral  margins  distinctly  but  rather  shallowly  concave.  Scutellum,  pleura, 
femora  and  veins  of  corium  granulate.  Abdomen  with  laterotergites  granulate. 

Colour  pale  yellowish  brown,  usually  with  darker  brown  mottling,  to  piceous  with  paler  markings. 
Antennal  segments  I  to  III  usually  pale  yellowish  brown  throughout,  infuscate  in  darkest  specimens,  III 
never  darkened  at  apex,  intercalary  segment  and  segment  IV  always  black.  Membrane  of  hemelytra 
colourless,  veins  white  with  short,  piceous  streaks. 

REMARKS.  Pseudophloeus  gestroi  was  described  from  a  single  male  taken  at  the  oasis  of  Jarabub  (Giarabub) 
in  Libya  by  Confalonieri.  In  his  original  description,  Bergevin  (1930:  32-33)  compared  it  with  falleni  and 
waltli  but  not  with  intermedius,  which  suggests  that  he  was  not  familiar  with  this  last  species.  His  description 
agrees  well  with  intermedius,  especially  in  the  'almost  smooth'  antennal  segment  III  and  in  the  shape  of  the 
pronotum,  which  is  usually  less  abruptly  widened  posteriorly  in  this  species  than  in  most  specimens  of 
falleni.  In  falleni,  antennal  segments  II  and  III  are  distinctly  more  strongly  granulate.  Bergevin  did  not 
mention  the  length  of  antennal  segment  I  in  relation  to  the  width  of  the  head,  which  is  greater  in 
intermedius  than  in  falleni.  A  drawing  of  the  holotype  of  P.  gestroi,  kindly  supplied  by  Dr  R.  Poggi, 
confirms  that  this  species  is  synonymous  with  the  former  rather  than  the  latter.  I  have  seen  specimens  of 
falleni  from  several  localities  each  in  Britain,  Spain  and  Greece  and  from  single  localities  in  Austria, 
Gibraltar  and  Tunisia  (Sbeitla).  The  specimen  of  intermedius  listed  below  from  Ethiopia  has  shorter 
appendages  than  the  others  examined.  Its  antennal  segment  I  was  found  to  be  almost  as  short  as  in  falleni.  It 
appears  to  have  suffered  some  damage  to  one  posterior  leg  and  to  the  abdominal  apex  during  development, 
which  suggests  that  it  may  not  be  typical  of  the  species  in  Ethiopia.  Dr  I.  M.  Kerzhner  (pers.  comm.) 
reports  two  specimens  from  Addis  Ababa  in  IZ. 

DISTRIBUTION.  Eremic.  Spain  and  Portugal  but  no  other  countries  in  Europe  to  the  north  of  the 


TRIBE  PSEUDOPHLOEINI  193 

Mediterranean  Sea;  North  Africa  south  to  Chad  and  Ethiopia;  Sinai;  Iraq;  Iran;  Turkey;  Turkmenia  (in 
part  fide  Stichel,  1962: 194). 

MATERIAL  EXAMINED 

Egypt:  1  $,  (holotype  of  angustus),  Sinai,  Ai'oun  Mousa,  ii.1899  (Autran)  (data  fide  Reuter,  1891) 
(ZMU). 

Chad:  2  cf,  Kanem  District,  N'Gouri,  viii.1958,  x-xi.1958  (Renoud)  (MRAC).  Sudan:  1  cf,  Um 
Enderaba,  2.x. 1927,  at  light  (Johnston)  (BMNH).  Ethiopia:  1  cf ,  Simien,  Ras  Degien,  below  the  pass, 
over  4300  m,  swept  from  Senecio  farinaceus  and  Helichrysum  citrispinum,  ll.xii.1952  (Scott)  (BMNH). 

Extralimital  material.  Spain:  4  $ ,  Cangas  (Champion).  Algeria:  3  cf ,  2  9 ,  Biskra  (Champion).  Tunisia: 
1  $,  Tozeur  (Champion);  2  $>,  Gafsa  (Champion);  2  $,  Sfax  (de  Vauloger).  Libya:  1  cf,  Cyrenaica, 
xi.1942.  Palestine:  1  cf ,  1  $,  Gaza  district,  Deir  El-Belah,  lO.v.1917, 14.V.1917  (Austen).  Iraq:  1  cf ,  2  $, 
Baghdad,  xii.1918,  27.xi.1918  (Harwood).  Iran:  2  $,  Shiraz,  Bushire,  1-2.V.1927  (Siyazov).  U.S.S.R.:  1 
Cf,  2  $,  Uzbekistan,  Termez,  24.V.1912,  5.V.1912,  25.V.1912  (Kirichenko);  2  cf,  1  $,  Tadzhikistan, 
Molotovabad  (=  Pyandzh),  5-7.iii.1944  (Kirichenko).  (All  BMNH.) 

MFLAStal 
Myla  Stal,  1866:  111.  Type-species:  Myla  nigrispina  Stal,  by  monotypy. 

Body  form  parallel-sided,  length  about  3  times  breadth,  sometimes  broader  (M.  lata),  sometimes 
somewhat  depressed  (M.  niokensis),  connexivum  narrow  to  broad. 

Head  about  as  long  as  broad,  weakly  granulate.  Antennifers  weakly  divergent,  their  outer  apical 
processes  porrect.  Antennae  with  segment  I  varying  in  length  from  about  0-50-1-25  times  width  of  head 
including  eyes,  almost  isodiametric  throughout  its  length,  weakly  granulate;  segment  III  about  1-25  times 
as  long  as  II,  both  segments  slender  and  weakly  granulate;  segment  IV  elongate,  cylindrical,  variable  in 
length  between  the  species  so  that  in  some  it  is  the  longest  and  in  others  the  shortest  segment,  its  specialized 
sensory  area  occupying  nine-tenths  of  its  length  or  more.  Bucculae  occupying  about  one-third  of  ventral 
length  of  head.  Rostrum  at  rest  reaching  posteriorly  to  base  of  metasternum. 

Pronotum  strongly  declivent  anteriorly,  disc  weakly  or  very  weakly  granulate;  lateral  margins  with  many 
small  granules  or  a  few  large  granules  or  tubercles;  posterolateral  angles  scarcely  to  very  strongly  produced 
anterolaterally  or  laterally,  bearing  obsolete  to  long,  slender  and  acute  posterolateral  spines;  posterior 
margin  weakly  convex,  smoothly  rounded,  sometimes  appearing  slightly  trilobed  but  without  distinct 
emarginations  or  projections.  Scutellum  equilateral  or  slightly  elongate,  its  apex  slightly  elevated  above 
clavi  at  rest,  usually  white;  lateral  margins,  especially  anteriorly,  slightly  elevated;  disc  flat,  weakly 
granulate-punctate.  Mesosternum  sulcate  throughout.  Metasternum  strongly  convex,  sulcate  only  anter- 
iorly. Metathoracic  scent-gland  peritreme  with  dorsal  ridge  reniform.  Corium  with  distal  margin  weakly  or 
very  weakly  concave,  its  apex  reaching  to  level  of  suture  between  laterotergites  V  and  VI  when  at  rest. 
Metathoracic  wing  with  antevannal  vein  present.  Anterior  and  intermediate  femora  usually  with  a  single, 
prominent,  subapical  granule  or  tubercle  beneath;  posterior  femur  with  well-developed  basal  tubercle  and 
subapically  beneath  with  three  major  spines,  of  which  the  first  (proximal)  one  is  very  small,  two  to  four 
small  granules  or  tubercles  between  the  two  larger  spines  and  an  apical  series  of  four  tubercles.  Posterior 
coxae  separated  by  a  distance  equal  to  approximately  half  the  width  of  a  coxa. 

Abdominal  sterna  III  to  VII  with  posterolateral  angles  acute,  weakly  to  strongly  produced,  lateral 
margins  of  these  sternites  finely  or  obsoletely  granulate.  Male  genital  capsule  posteriorly  emarginate,  the 
emargination  with  a  central  cusp;  parameres  with  blades  flat,  their  apices  not  occluding  emargination  of 
capsule  (Fig.  96),  proximal  tooth  of  paramere  simple,  apical  tooth  long,  upcurved.  Phallotheca  (Figs 
111-113)  comprising  a  single  dorsal  and  a  single  ventral  sclerite,  both  broad,  each  with  a  narrow,  median, 
posterior  projection,  ventral  sclerite  proximally  produced  dorsad  on  each  side  to  meet  dorsal  sclerite. 
Conjunctiva  (Figs  111-114)  with  dorsomedian  lobe  membranous,  transverse,  its  lateral  angles  prominent; 
distal  dorsolateral  lobes  membranous,  arising  close  together  over  ejaculatory  reservoir,  finger- like, 
reflexed  anteriad;  apical  ventral  lobes  large,  membranous,  paired;  distal  ventrolateral  lobes  deeply  bifid, 
wings  of  ejaculatory  reservoir  complex  extending  into  the  dorsal,  descending  arm  of  each  lobe,  ventral, 
anteriorly  directed  arm  of  each  lobe  membranous  and  without  sclerotized  support;  ventral  lobes  lightly 
sclerotized,  wide  apart,  narrow,  anteriorly  directed  and  slightly  upcurved;  base  of  vesica  without 
protective  sclerites.  Sclerites  of  wall  of  gynatrium  in  female  each  comprising  a  curved  plate  parallel  to  rami 
bearing  a  longer,  ventral,  mesal  and  a  shorter,  dorsal,  lateral  process,  both  directed  posteriorly. 
Spermatheca  (Figs  115,  116)  with  bulb  lunate,  duct  adjacent  to  bulb  tightly  convoluted,  connected  with 
gynatrium  by  an  S-shaped  portion  of  the  duct. 

Main  pubescence  of  body,  hemelytra,  femora  and  first  antennal  segment  consisting  of  short,  semidecum- 


194  W.  R.  DOLLING 

bent  to  adpressed,  white,  flattened,  scale-like  hairs.  Coloration,  probably  of  all  species,  variable,  basically 
grey-brown  with  more  or  less  of  a  yellowish  or  reddish  suffusion. 

REMARKS.  This  genus  differs  from  all  others  in  the  tribe  in  the  form  of  the  parameres,  of  which  the  distal 
tooth  is  long  and  sharply  upcurved  at  right  angles  to  the  blade  and  the  apices  do  not  fill  the  posterior 
emargination  of  the  genital  capsule.  The  form  of  the  phallotheca  is  also  unique  within  the  tribe.  The 
scale-like  pubescence  is  sufficient  to  distinguish  Myla  species  from  all  other  African  Pseudophloeinae 
except  Paramyla  australis.  Myla  species  are  sometimes  mistaken  for  members  of  the  tribe  Clavigrallini 
which,  however,  lack  the  tubercle  that  is  present  in  Myla  at  the  base  of  the  posterior  femur.  Eleven  species, 
all  African,  are  accepted  here  as  members  of  the  genus.  For  M.  concolor  Dohrn  and  M.  cornuta  Hsiao,  see 
Pseudomyla;  for  M.  suspecta  Schouteden,  see  Paramyla.  M.  schnelli  Villiers  was  transferred  to  Clavigralla 
by  Dolling  (19790:  34). 

DISTRIBUTION.  Africa  south  of  the  Sahara,  including  Ethiopia.  Not  so  far  recorded  from  any  islands  or  from 
Arabia. 

Key  to  species 

1  Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes  0-84  or  less;  in  doubtful 

cases  width  of  pronotum  across  apices  of  posterolateral  spines  divided  by  width  of  head  2-22 

or  less 2 

-  Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes  0  •  85  or  more ;  in  doubtful 

cases  width  of  pronotum  across  apices  of  posterolateral  spines  divided  by  width  of  head  more 
than2-30 4 

2  Length  of  antennal  segment  I  divided  by  width  of  head  less  than  0-54.  (East  Africa,  rare) 

d/spar(p.  198) 

-  Length  of  antennal  segment  I  divided  by  width  of  head  more  than  0-54 3 

3  Pronotum  (Fig.  85)  with  posterolateral  spines  very  short.  (Mountains  of  East  Africa)ni'olcensis(p.  201) 

-  Pronotum  (Fig.  80)  with  posterolateral  spines  longer.  (Cameroun) onceroma  (p.  203) 

4  Pronotum  (Fig.  89)  with  posterolateral  spines  short,  laterally  directed;  tibiae  (Fig.  95)  with 

alternate  dark  and  light  annuli.  (East  Africa,  rare) /afa (p.  200) 

-  Pronotum  (Figs  79,  81-84,  86-88)  with  posterolateral  spines  longer,  directed  anterolaterally; 

tibiae  stramineous,  unicolorous 5 

5  Pubescence  of  body  (Fig.  98)  long,  individual  scale-like  hairs  of  thorax  longer  than  distance 

between  their  insertions.  (West  and  Central  Africa) gracilis  (p.  204) 

-  Pubescence  of  body  (Fig.  97)  short,  individual  scale-like  hairs  of  thorax  shorter  than  distance 

between  their  insertions 6 

6  Ocelli  (Fig.  81)  very  large,  distance  between  ocellus  and  eye  less  than  half  diameter  of  ocellus. 

(East  Africa) somalica  (p.  200) 

-  Ocelli  (Figs  79, 82)  smaller,  distance  between  ocellus  and  eye  equal  to  or  greater  than  diameter 

of  ocellus 7 

7  Head  (Fig.  82)  elongate,  apically  acute,  its  midline  fuscous;  eyes  small.  (Widespread  in  Africa) 

microphthalma(p.  202) 

-  Head  (Fig.  79)  short ,  apically  obtuse ,  its  midline  not  fuscous ;  eyes  larger 8 

8  Lateral  margins  of  pronotum  (Fig.  86)  bearing  large,  prominent  granules.  (Southern  Africa, 

rare) granula  (p.  198) 

Lateral  margins  of  pronotum  (Figs  79, 84, 87, 88)  with  smaller  granules 9 

9  Posterolateral  angles  of  pronotum  and  abdominal  sternites  (Fig.  92)  less  prominent.  (Ethiopia) 

abyssinica(p.  197) 
Posterolateral  angles  of  pronotum  (Figs  79,  84,  87,  88)  and  abdominal  sternites  (Fig.  91)  more 

prominent 10 

10    Paramere  (Figs  99, 100)  with  apical  tooth  projecting  beyond  apex  of  shaft;  second  valvula  (Fig. 

119)  apically  rounded.  (West  and  Central  Africa) hoploxys(p.  194) 

Paramere  (Fig.  104)  with  apical  tooth  not  projecting  beyond  apex  of  shaft;  second  valvula  (Fig. 

120)  obliquely  truncate,  with  apex  acute.  (Central,  eastern  and  southern  Africa) . . .      calida(p.  197) 

Myla  hoploxys  (Dallas) 
(Figs  79,  84,  91,  96,  99, 100, 119) 
Clavigralla!  hoploxys  Dallas,  1852:  515.  Holotype  cf ,  GAMBIA  (BMNH)  [examined]. 


TRIBE  PSEUDOPHLOEINI 


195 


79 


Figs  79-94  My  la  species.  79-82,  dorsal  view  of  head  and  pronotum  of  (79)  hoploxys;  (80)  onceroma;  (81) 
somalica;  (82)  microphthalma.  83-89,  dorsal  view  of  pronotum  of  (83)  gracilis ,  lectotype;  (84)  hoploxys; 
(85)  niokensis;  (86)  granula;  (87)  calida  from  Zaire:  Mpese;  (88)  calida,  holotype  from  Uganda;  (89) 
lata,  holotype.  90-94,  ventral  view  of  abdominal  margin  of  (90)  niokensis;  (91)  hoploxys;  (92)  abyssinica; 
(93)  lata,  holotype;  (94)  gracilis . 


196  W.  R.  DOLLING 

Myla  nigrispina  Stal,  1866:  111.  LECTOTYPE  $,  'GUINEA'  (NMV),  here  designated  [examined].  Syn.  n. 

Myla  hoploxys  (Dallas)  Stal,  1873:  84. 

Myla  hoploxys  nigrispina  Stal;  Linnavuori,  1971: 176. 

Length:  cf,  9-8-11-7 mm;  $,9-7-12-0 mm. 

Head  (Fig.  79)  slightly  longer  than  wide,  broadly  rounded  anteriorly.  Length  of  antennal  segment  I 
divided  by  width  of  head  including  eyes  in  male  0  •  88-1  •  16 ,  in  female  0  •  89-1  •  09 .  Ratio  of  lengths  of  antennal 
segments  in  male  about  l-00:0-80:l-03:0-96,  in  female  about  l-00:0-84:l-04:0-85.  Length  of  rostral 
segment  I  divided  by  width  of  head  including  eyes  in  both  sexes  0-69-0-81,  ratio  of  lengths  of  rostral 
segments  about  1 -00:0-84:0-43 :0-64. 

Pronotum  (Figs  79,  84)  with  posterolateral  angles  produced  anterolaterally  and  terminating  in  short 
spines,  lateral  margins  very  weakly  granulate.  Width  across  apices  of  posterolateral  spines  divided  by  width 
of  head  including  eyes  in  male  2-14-2-84,  in  female  2-20-2-92.  Length  of  posterior  tibia  divided  by  that  of 
posterior  femur  0-89-1-06. 

Abdomen  rather  narrow,  its  lateral  margin  (Fig.  91)  weakly  convex,  posterolateral  angles  of  sternites 
III- VII  projecting  as  fairly  short,  narrow,  posteriorly  directed  spines;  abdominal  laterotergites  narrow, 
laterotergite  V  usually  at  least  twice  as  long  as  wide,  rarely  slightly  less.  Paramere  (Figs  96,  99, 100)  with 
apical  tooth  long,  upcurved,  its  posterior  margin  projecting  distinctly  beyond  apex  of  shaft.  Aedeagus  very 
similar  in  all  respects  to  that  of  M.  calida,  q.v.  Female  with  second  valvula  of  ovipositor  (Fig.  119) 
terminating  in  an  apically  rounded  lobe. 

Pubescence  of  head,  all  exposed  parts  of  thorax,  abdominal  laterotergites,  antennal  segment  I,  femora, 
clavus  and  corium  composed  of  short,  flattened,  scale-like,  decumbent  or  adpressed,  white  hairs,  each  hair 
much  shorter  than  the  distance  between  its  own  insertion  and  that  of  its  nearest  neighbour,  each  insertion 
borne  on  a  large  granule  on  the  head  and  a  small  granule  elsewhere,  granules  of  thoracic  pleurites, 
posterior  lobe  of  pronotum,  scutellum  and  hemelytra  each  positioned  on  the  anterior  (or  proximal  on 
hemelytra)  border  of  a  large  puncture.  Pubescence  of  disc  of  abdominal  venter  suberect,  not  scale-like, 
grading  laterally  into  white,  scale-like  type.  Pubescence  of  antennal  segments  II  to  IV,  tibiae  and  tarsi 
suberect,  rather  short,  not  scale-like. 

General  coloration  of  body  and  appendages  greyish  yellow,  more  or  less  heavily  suffused  red.  Antennal 
segments  II  and  III,  tibiae  and  tarsi  stramineous.  Apex  of  rostral  segment  IV,  posterolateral  angles  and 
spines  of  pronotum  and  often  a  line  along  lateral  margins  of  pronotum  piceous.  Membrane  of  hemelytra 
whitish  hyaline,  its  veins  brown  with  frequent  whitish  hyaline  interruptions.  Elevated  apex  of  scutellum 
and  dorsal  ridge  of  metathoracic  peritreme  white,  the  latter  often  margined  piceous. 

REMARKS.  This  and  the  next  four  species  form  a  group  of  closely  related  taxa  which  are  probably  all  rather 
variable  in  pronotum  shape,  size  and  coloration.  Examination  of  the  genitalia  is  the  only  completely 
reliable  way  of  distinguishing  between  this  species  and  M.  calida,  with  which  it  is  partly  sympatric. 
Although  the  genitalia  of  the  female  lectotype  of  M.  nigrispina  were  not  dissected  out,  the  specimen 
corresponds  in  every  other  way  with  the  West  African  species  recognized  here  as  M.  hoploxys.  No  other 
species  of  this  complex  was  detected  in  the  area  of  the  type-locality  of  M.  nigrispina  ('Guinea',  =  West 
Africa).  There  is  no  justification  for  Linnavuori's  (1971:  175,  176)  recognition  of  two,  largely  sympatric 
subspecies  distinguished  by  size,  colour  and  some  slight,  probably  allometric  structural  features.  Large, 
red  individuals  with  long  posterolateral  pronotal  spines  predominate  in  the  western  part  of  the  range  of  the 
species  and  are  rare  in  the  eastern  part.  This  species  includes  cowpea  (Vigna)  in  its  diet  and,  according  to 
Professor  A.  E.  Akingbohungbe  (pers.  comm.),  it  is  approaching  the  pest  status  of  the  Clavigralla  species 
on  this  crop. 

DISTRIBUTION.  Proven  distribution  covers  West  Africa  from  Senegal  and  Gambia  eastwards  to  Zaire  and 
Angola.  Linnavuori  (1978:  36)  records  a  single  specimen  from  Sudan. 

MATERIAL  EXAMINED 

Gambia:  1  cf  (holotypeof  hoploxys),  (no  other  data)  (BMNH).  'Guinea':  1  $  (lectotype  of  nigrispina) 
(coll.  Signoret),  (NMV). 

Gambia:  1  cf  (no  further  locality  data;  pin  and  locality  label  identical  with  those  of  holotype  of  C. 
hoploxys  but  also  with  label:  'Saunders  Coll.');  1  $ ,  3.iii.l911  (Simpson)  (BMNH).  Senegal:  1  cf ,  Bambey, 
6.V.1943  (Risbec)  (BMNH);  1  Cf,  2  $,  M'Bambey,  28.x-6.xi.  1939  (Risbec)  (MRAC).  Ivory  Coast:  4  cf , 
Lamto,  7.V.1963,  14.V.1963,  28.V.1963,  18.vi.1963;  3  $,  Lamto  (Toumodi),  18. ii. 1964,  24.vii.1964;  1  $>, 
Bouake,  21.vii.1962  (probably  all  leg.  Gillon)  (BMNH);  2  cf,  Bouake,  ii.1968  (Schmitz)  (MRAC). 
Ghana:  1  $,  Tafo,  8.xi.l965  (Leston);  1  cf,  Labadi,  12.ix.1965  (Lestori)  (BMNH);  1  cf,  Namgua, 
30.ii.1964, oncowpeas;  1  cf ,  1  $,  Sokode, 7.xi.l967 (Leston);  1  $,  (locality illegible),  8.xii.l965  (Leston); 
1  $ ,  Legon,  22.ix.1968,  in  u.v.  trap  (Kumar);  1  $,  Abouadi,  7.xi.l976  (Leston)  (UG).  Mali:  1  cf ,  40  miles 


TRIBE  PSEUDOPHLOEINI  197 

(64  km)  SW.  of  San,  300  m,  26.viii.1966  (Ross,  Lorenzen)  (CAS).  Nigeria:  1  cf,  Ibadan,  30.xi.1976 
(Deeming)  (IAR);  1  cf,  He-Ife,  5.xii.l970  (Medler);  1  $,  Ile-Ife,  14.ii.1974  (Akingbohungbe);  1  $, 
Badeggi,  19.iii.1972  (Medler)  (He  University);  1  $,  Azare,  1928-1929  (Lloyd);  1  cf ,  1  $,  Zaria,  Samaru, 
9.viii.l966,  21. ix. 1967  (Deeming);  1  $,  Zaria,  Samaru,  14. ix. 1966,  on  cowpea  (Vigna)  (Deeming);  1  9, 
Samaru,  ll.iii.1955,  at  grass  roots  (Emsley);  4  cf ,  1  $,  Samaru,  7-14. vii.  1970,  28.viii.1970,  8-15.ix.1970 
(Ward);  1  $,  Ilorin,  19.iii.1955,  in  dry  humus  (Emsley);  1  $,  Jos,  9.X.1955  (Bechyne);  1  cf ,  Ibadan,  DFR 
nursery,  yellow  tray  trap,  30. ix. 1963  (White);  1  cf ,  Enugu,  22.x. 1955  (Bechyne);  1  $,  Lagos  'hinterland' 
(Rowland)  (BMNH);  1  cf ,  1  $,  Jos  province,  7-26.iv.1963  (Meussen,  Bouquiaux)  (MRAC).  Zaire:  1  $, 
Bambesa,  22.U938  (Vrydagh);  1  $,  Katanga,  Nyonga  (?  =  Nyanga),  v.1925  (de  Witte);  1  $,  Wombali, 
vii.1919  (Vanderijst);  1  cf ,  2  $,  Lulua,  Kapanga,  iii.1932,  iii.1933  (Overlaet)  (MRAC).  Angola:  1  cf ,  Joao 
de  Almeida,  29.iii.1972  (BMNH  Expedition)  (BMNH).  'Tropical  Africa':  2  cf  (UM). 

Myla  abyssinica  Linnavuori 
(Figs  92,  97, 102) 

Myla  abyssinica  Linnavuori,  1971:  175.  Lectotype  cf ,  ETHIOPIA:  near  Nazareth,  20-21.vi.1963  (Linna- 
vuori) (R.  Linnavuori  coll.),  designated  by  Linnavuori,  1978:  36  [not  examined]. 

Length:  cf ,  9-9-10-1  mm;  $  unknown. 

Very  similar  to  M.  hoploxys.  Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes 
0-88-0-93.  Ratio  of  lengths  of  antennal  segments  about  1-00:0-80: 1-05: 1-04.  Length  of  rostral  segment  I 
divided  by  width  of  head  including  eyes  0-69-0-75.  Ratio  of  lengths  of  rostral  segments  about 
1 -00:0-84:0-45 :0-64.  Pronotum  with  posterolateral  spines  as  short  as  in  the  shortest-spined  examples  of  M. 
hoploxys;  width  across  apices  of  spines  divided  by  width  of  head  including  eyes  2-16-2-21.  Connexivum 
narrow,  laterotergite  V  about  2-5  times  as  long  as  wide.  Posterolateral  spines  of  sternites  III  to  VII  short 
(Fig.  92).  Length  of  posterior  tibia  divided  by  length  of  posterior  femur  0-96.  Paramere  (Fig.  102)  with 
apical  tooth  relatively  shorter  than  that  of  M.  hoploxys. 

REMARKS.  This  species  was  described  by  Linnavuori  (1971:  175)  in  a  dichotomous  key.  Subsequently,  the 
same  author  (Linnavuori,  1978:  21,  36, 103,  figs  16f,  16g,  69d)  provided  a  fuller  description,  with  figures, 
suggesting  at  the  same  time  that  it  might  be  only  a  race  of  M.  hoploxys.  In  his  1978  publication,  he 
designated  his  unique  specimen  as  holotype.  As  he  already  validly  described  the  species  in  1971,  without 
designating  a  type-specimen,  his  1978  action  must  be  regarded  as  a  lectotype  designation.  In  view  of  the 
small  differences  separating  species  in  this  genus  it  seems  appropriate  to  accord  full  specific  status  to  this 
taxon,  especially  as  the  two  specimens  examined  here  are  very  similar  to  the  one  described  by  Linnavuori. 
The  strong  reddish  suffusion  noted  by  Linnavuori  (1978)  in  the  type-specimen  is  not  apparent  in  the  two 
specimens  I  have  seen. 

DISTRIBUTION.  Endemic  to  Ethiopia  on  the  available  evidence. 

MATERIAL  EXAMINED 

Ethiopia:  1  cf ,  Djoudjou  Abayi,  Didessa  R.,  Goma,  2000  ft  (600  m),  lO.v.1905  (Zaphiro)  (BMNH);  1 
Cf,  Gemu-Gofa  Prov.,  45  km  N.  of  Demika,  near  Jinka  road,  1460  m,  30. iv.  1974  (de  Rougemont) 
(MRAC). 

Myla  calida  sp.  n. 

(Figs  87,  88, 101, 111-114, 120) 

Length:  cf ,  9-0-10-6  mm;  $ ,  9-0-10-3  mm. 

Very  similar  to  M.  hoploxys.  Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes  in 
male  and  female  0-92-1-05.  Ratio  of  lengths  of  antennal  segments  in  male  about  l-00:0-75:0-95:0-94,  in 
female  about  l-00:0-80:0-95:0-82.  Length  of  rostral  segment  I  divided  by  width  of  head  including  eyes  in 
male  0-66-0-75,  in  female  0-69-0-79.  Ratio  of  lengths  of  rostral  segments  in  male  about 
l-00:0-85:0-43:0-64,  in  female  about  1-00:0-85:0-42:0-61. 

Pronotal  shape  variable  (Figs  87,  88),  posterolateral  spines  on  average  more  strongly  divergent  than 
those  of  M .  hoploxys;  width  across  apices  of  posterolateral  spines  divided  by  width  of  head  including  eyes 
in  male  2-30-2-85,  in  female  2-31-2-62.  Granulation  of  lateral  margins  of  pronotum  slightly  more 
pronounced  than  in  M.  hoploxys.  Length  of  posterior  tibia  divided  by  length  of  posterior  femur  0-89-0-95. 

Connexivum  and  abdominal  spines  as  in  M.  hoploxys.  Paramere  (Fig.  101)  with  apical  tooth  shorter  than 
that  of  M.  hoploxys,  its  posterior  margin  not  or  scarcely  projecting  posteriad  of  apex  of  shaft.  Aedeagus 


198  W.  R.  DOLLING 

(Figs  111-114)  of  the  form  usual  in  the  genus.  Ovipositor  with  second  valvula  (Fig.  120)  obliquely  truncate, 
apex  acute  and  slightly  upturned. 

Coloration  on  average  more  yellowish  than  greyish,  never  as  deeply  suffused  red  as  the  most  extreme 
examples  of  M.  hoploxys. 

REMARKS.  The  second  valvula  of  this  species  is  very  distinctive.  The  shape  of  the  paramere  is  rather  variable 
but  it  is  never  so  long  as  that  of  M.  hoploxys,  nor  is  its  apical  tooth  so  long  or  so  prominent.  The  shape  of  the 
pronotum  is  also  variable  (compare  Figs  87, 88),  as  is  the  colour.  In  many  individuals  there  is  a  conspicuous 
black  line  along  the  lateral  margins  of  the  pronotum. 

DISTRIBUTION.  Eastern,  central  and  southern  Africa.  Absent  from  West  Africa.  Northern  limit  unknown. 

MATERIAL  EXAMINED 

Holotype  cf ,  Uganda:  W.  shores  of  Vic.  Nyanza,  Buddu,  3700  ft  (1100  m),  19-25.ix.1911  (5.  A.  Neave) 
(BMNH). 

Paratypes.  Zaire:  1  cf,  1  $,  Ngowa,  5.vi.l939,  16.vi.1939  (Mertens);  1  cf,  Kwango,  Ngowa,  3.X.1937 
(Mertens);  1  C? ,  1  ?,  Ngowa-Kwango,  no  date  and  17. ii. 1938  (Mertens);  1  cf ,  Kibangula,  1957  (Henry);  1 
Cf,  2  $,  Mpese,  7-26.vi.1937  (Cooreman)  (IRSNB);  1  tf,  Sankuru,  M'Pemba  Zeo  (Gandajika), 
17.vii.1958  (Marechaf);  1  cf ,  Kwango,  Popokabaka,  v.1952  (Pierquin);  1  cT,  Kasai,  Ilebo  (Thery);  1  cf , 
Bokala,  20.V.1915  (Mayne);  1  cf ,  Tolo,  early  xii.1913  (Maes);  1  cf ,  Kivu,  Kavimvira  (Uvira),  vi.1955,  at 
light  (Marlier);  1  cf,  Kinshasa  (Tynant);  1  cf,  Lita,  25.V.1912  (Christy);  1  cf,  'Dans  le  ebenal.  Post 
Telegraphiq  au  dessous  de  Block  river,  22  Mai  1909.  Herbes  et  arbustes  a  3-4  au  dessus  du  fleuve,  endroit 
non  boise,  1  a  17  heures'  (Voyage  de  S.A.R.  le  Prince  Albert);  1  $ ,  Kisantu,  1925  (Varderijst);  2  $ ,  Lulua, 
Kapanga,  i.1933,  v.1933  (Overlaet);  1  $,  Sankuru,  M'Pemba  Zeo  (Gandajika),  28.iv.1960  (Marechaf);  1 
$,  Mayidi,  1942  (van  Eyen);  1  ex.  without  abdomen,  Bas-congo,  Lemfu,  vi.1945  (de  Beir)  (MRAC);  6  cf , 
13  $,  Faradje,  29°40'E,  3°40'N  (Lang,  Chapin)  (AMNH).  Zambia:  1  cf,  1  $,  High  Plateau,  L. 
Tanganyika,  4500  ft  (1350  m),  1 8-2 l.viii.  1908  (Neave)  (UM).  Angola:  1  cf  (other  data  illegible)  (BMNH). 
Zimbabwe:  1  cf ,  Salisbury,  v.1963  (NMB);  1  $ ,  Umtali  (BMNH).  South  Africa:  1  cf ,  Durban,  1902  (Muir) 
(BMNH). 

/V/v/a  d/sparsp.  n. 

(Fig.  104) 

Length:  cf ,  9-6  mm;  $ ,  unknown. 

Resembles  M.  hoploxys  in  most  respects,  including  the  shape  of  the  paramere  (Fig.  104),  which 
distinguishes  it  from  M.  calida.  Lateral  margins  of  pronotum  rather  more  strongly  granulate  than  those  of 
M.  calida  and  posterolateral  spines  of  pronotum  short.  Length  of  antennal  segment  I  divided  by  width  of 
head  including  eyes  0-53.  Ratio  of  lengths  of  antennal  segments  as  1-00:0-82:1-06:1-18.  Width  of  pronotum 
across  apices  of  posterolateral  spines  divided  by  width  of  head  including  eyes  2-22.  Length  of  posterior  tibia 
divided  by  that  of  posterior  femur  0-92. 

REMARKS.  Perhaps  this  is  an  outlying  representative  of  M.  hoploxys  but  it  differs  from  that  species  and  from 
M.  calida  in  the  relative  lengths  of  the  antennal  segments,  of  which  I  and  II  are  very  short  and  III  and  IV 
longer  with  respect  to  I. 

DISTRIBUTION.  Zambia. 

MATERIAL  EXAMINED 
Holotype  cf ,  Zambia:  L.  Bangweolo,  Chirui  Island,  3800  ft  (1140  m),  5-7. vii. 1908  (Neave)  (UM). 

Myla  granula  sp.  n. 

(Figs  86, 121) 

Length:  cf ,  unknown;  $,  10-2-10-4  mm. 

Resembles  M.  hoploxys  and  M.  calida  in  general  form.  Length  of  antennal  segment  I  divided  by  width  of 
head  including  eyes  0-85-0-93.  Ratio  of  lengths  of  antennal  segments  in  holotype  as  1-00:0- 84: 1-00:0-95,  in 
paratype  as  1-00: 0-82: 0-92:0-91.  Length  of  rostral  segment  I  divided  by  width  of  head  including  eyes 
0-66-^0-69.  Ratio  of  lengths  of  rostral  segments  in  holotype  as  1-00:0-87:0-40:0-60,  in  paratype  as 
1-00:0-94:0-44: 0-67.  Pronotum  (Fig.  86)  with  posterolateral  angles  produced  laterally  almost  perpendicu- 
lar to  axis  of  body,  posterolateral  spines  very  short,  width  across  apices  of  spines  divided  by  width  of  head 
including  eyes  2-34-2-37;  lateral  margins  bearing  very  large  granules.  Length  of  posterior  tibia  divided  by 


TRIBE  PSEUDOPHLOEINI 


199 


95 


97 


98 


99 


101 


102 


103 


104 


106 


107 


108 


109 


Figs  95-110  My  la  species.  95,  lata,  posterior  tibia.  96,  hoploxys,  posterior  view  of  male  genital  capsule 
with  parameres.  97,  98,  detail  of  pubescence  of  left  metapleuron  of  (97)  abyssinica;  (98)  gracilis.  99, 
hoploxys,  dorsomedial  view  of  left  paramere.  100,  hoploxys,  lateral  view  of  same.  101, 102,  dorsomedial 
view  of  left  paramere  of  (101)  calida;  (102)  abyssinica.  103,  somalica,  dorsomedial  view  of  right 
paramere,  104-110,  dorsomedial  view  of  left  paramere  of  (104)  dispar;  (105)  microphthalma;  (106) 
niokensis;  (107)  lata;  (108)  gracilis  from  Zaire:  Ngowa;  (109, 110)  gracilis,  two  specimens  from  Zaire: 
Faradje. 


200  W.  R.  DOLLING 

length  of  posterior  femur  0-95-0-99.  Ovipositor  with  second  valvula  (Fig.  121)  symmetrically  tapering 
towards  narrowly  rounded  apex. 

REMARKS.  This  species  differs  from  M.  hoploxys  and  M.  calida  mainly  in  the  shape  and  granulosity  of 
pronotum.  The  second  valvulae  of  the  ovipositor  are  characteristic;  males  are  unknown. 

DISTRIBUTION.  South  Africa. 

MATERIAL  EXAMINED 

Holotype  $,  South  Africa:  Natal,  Malvern,  ll.vi.1897  (BMNH). 

Paratype.  South  Africa:  1  $ ,  Natal,  Lewombo  Mission,  31°40'E,  28°35'S,  15.x.  1977, 365  m,  under  brick 
pile(/?eave//)(NMP). 

Myla  somalica  Linnavuori 
(Figs  81, 103) 

Myla  somalica  Linnavuori,  1982:  14-16.  Holotype  cf ,  SOMALI  REPUBLIC:  Giohar,  18-20. iv. 1968  (Spedi- 
zione  Biologica  in  Somali  del  Centra  di  Studio  per  la  Faunistica  ed  Ecologia  Tropicali  del  Consiglio 
Nazionale  delle  Ricerche)  (Museo  Zoologico  delPUniversita  di  Firenze)  [not  examined]. 

Length:  cf,  10-5-11-5  mm;  $  unknown. 

An  elongate  species  related  to  M.  calida.  Ocelli  very  large  (Fig.  81).  Length  of  antennal  segment  I 
divided  by  width  of  head  including  eyes  0-85-0-99.  Ratio  of  lengths  of  antennal  segments  about 
1-00:0-82:0-97: 1-08.  Length  of  rostral  segment  I  divided  by  width  of  head  including  eyes  0-70-0-71.  Ratio 
of  lengths  of  rostral  segments  about  1  -00:0-87:0-42:0-66. 

Pronotum  (Fig.  81)  with  posterolateral  angles  shortly  produced  laterally,  with  slight  anteriad  inclination; 
width  across  apices  of  posterolateral  spines  divided  by  width  of  head  including  eyes  2-31-2-44.  Posterior 
tibia  long,  its  length  divided  by  that  of  posterior  femur  0-96-1-03. 

Paramere  (Fig.  103)  with  shaft  long,  as  in  M.  hoploxys,  teeth  of  blade  short,  as  in  M.  calida. 

REMARKS.  This  species  is  readily  distinguished  from  all  others  of  the  genus  by  its  enormous  ocelli.  The 
paratype  was  received  after  the  above  measurements  were  taken.  It  is  9-8  mm  long,  its  first  antennal 
segment  is  0-90  times  the  width  of  the  head  and  the  antennal  ratio  is  1-00:0-79: 1-07: 1-08.  The  paramere  is 
indistinguishable  from  that  of  the  inland  specimens  with  which  it  seems  to  be  conspecific  despite  the 
difference  in  habitat. 

DISTRIBUTION.  Highlands  of  eastern  central  Africa  to  the  Somali  coast. 

MATERIAL  EXAMINED 

Somali  Republic:  1  cf  (paratype).  Giohar,  18-20.iv.1968  (L.  Simonettd)  (R.  Linnavuori  coll.).  Kenya:  1 
Cf ,  Nakuru,  viii.1912  (van  Someren)  (BMNH).  Uganda:  1  ex.  without  abdomen  (probably  cf),  Chisinga, 
viii.1911  (Marshall)  (BMNH).  Zaire:  3  cf,  Pare  National  Albert,  Camp  Ruindi  (or  Rwindi),  1000  m, 
20-28.xi.1934  (de  Witte);  2  cf ,  P.N.A.,  Cp  Ruindi,  13.ix.1932  (Burgeon)  (MRAC). 

Myla  lata  sp.  n. 

(Figs  89,  93,  95, 107) 

Length:  cf ,  9-8-10-8  mm;  $  unknown. 

A  very  broad-bodied  species.  Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes  in 
holotype  0-88.  Remainder  of  antennae  in  holotype  and  whole  of  antennae  in  paratype  missing.  Length  of 
rostral  segment  I  divided  by  width  of  head  including  eyes  in  paratype  0-70;  ratio  of  lengths  of  its  rostral 
segments  as  l-00:0-80:0-45:0-63;  rostrum  not  visible  in  holotype  mount. 

Pronotum  (Fig.  89)  very  broad,  lateral  margins  strongly  granulate,  posterolateral  angles  weakly 
produced,  terminating  in  small,  laterally  directed  spines;  width  across  apices  of  posterolateral  spines 
divided  by  width  of  head  including  eyes  2-52-2-56. 

Abdomen  broad,  its  lateral  margins  (Fig.  93)  strongly  convex,  posterolateral  angles  of  sternites  III  to  VII 
produced  into  broad,  triangular  spines.  Connexivum  broad,  laterotergite  V  about  five-sixths  as  broad  as 
long.  Paramere  (Fig.  107)  with  both  teeth  of  blade  on  a  common  stem. 

Coloration  ferruginous;  all  tibiae  stramineous  with  basal,  median  and  apical  annuli  piceous  (Fig.  95). 

REMARKS.  This  species  is  very  distinctive  by  reason  of  its  great  breadth.  It  is  the  only  species  of  the  genus  in 
which  the  tibiae  are  conspicuously  bicolorous. 


TRIBE  PSEUDOPHLOEINI  201 

DISTRIBUTION.  Central  Africa. 

MATERIAL  EXAMINED 

Holotype  cf ,  Zambia:  N.  Lake  Bangweolo,  4200  ft  [1260  m],  Luwingu,  21.vii.1908  (Neave)  (UM). 
Paratype.  Zaire:  1  cf ,  Katanga,  Lufira  River,  3500  ft  [1050  m],  10.ix.1907  (Neave)  (BMNH). 

Myla  niokensis  Schouteden 
(Figs  85,  90, 106) 

Myla  niokensis  Schouteden,  1938:  294.  LECTOTYPE  cf ,  ZAIRE  (MRAC),  here  designated  [examined]. 
Myla  (Paramyla)  niokensis  Schouteden;  Linnavuori,  1971: 178. 

Length:  cf,  8-2-9-3  mm;  $,8-7-9-7 mm. 

Head  similar  in  shape  to  that  of  M.  hoploxys.  Antennae,  especially  segment  I,  short.  Length  of  antennal 
segment  I  divided  by  width  of  head  including  eyes  in  male  0-56-0-74,  in  female  0-55-0-69.  Ratio  of  lengths 
of  antennal  segments  in  male  about  l-00:0-95:l-14:l-48,  in  female  about  l-00:l-04:l-09:l-33.  Length  of 
rostral  segment  I  divided  by  width  of  head  including  eyes  in  both  sexes  0-64-0-75 .  Ratio  of  lengths  of  rostral 
segments  about  l-00:0-85:0-44:0-64. 

Pronotum  (Fig.  85)  with  posterolateral  angles  produced  forward,  their  apices  directed  laterally  but  very 
weakly  produced  in  that  direction,  terminating  in  very  short  spines,  margins  behind  the  produced  angles 
concave,  diverging  posteriorly  so  that  in  most  cases  greatest  width  of  pronotum  is  not  across  apices  of 
posterolateral  spines  but  across  posterior  lobe;  width  across  apices  of  posterolateral  spines  divided  by 
width  of  head  including  eyes  in  male  1-88-2-16,  in  female  2-00-2-22.  Length  of  posterior  tibia  divided  by 
length  of  posterior  femur  in  both  sexes  0-94-1-06. 

Abdomen  with  lateral  margin  (Fig.  90)  moderately  convex;  posterolateral  angles  of  sternites  III  to  VI 
weakly  prominent.  Connexivum  moderately  broad,  laterotergite  V  about  1-7  times  as  long  as  wide. 
Paramere  (Fig.  106)  very  similar  to  that  of  M.  hoploxys. 

Colour  yellow-brown  with  punctures  of  pronotum  and  hemelytra  and  often  parts  of  underside  and 
posterior  femora  piceous  or  black;  body  and  appendages  often  with  a  reddish  tinge.  The  darkest  species  of 
the  genus.  Pubescence  as  in  M.  hoploxys.  Granules  of  lateral  margins  of  pronotum  about  as  prominent  as 
those  of  M.  hoploxys. 

REMARKS.  This  species  may  easily  be  recognized  by  the  form  of  the  pronotum,  except  that  it  might  be 
confused  with  M.  abyssinica  from  which  it  differs  in  that  its  antennal  segment  I  is  much  shorter  and  segment 
IV  is  very  long  compared  with  I.  Specimens  from  Zaire  have  the  apical  tooth  of  the  paramere  slightly 
shorter  than  it  is  in  those  from  Ethiopia  and  Kenya.  Ethiopian  specimens  differ  from  the  others  in  having  a 
slightly  longer  rostral  segment  IV.  Antennal  segment  I  is  shortest,  both  in  proportion  to  head  width  and  in 
proportion  to  the  other  segments,  in  Kenyan  specimens  and  longest  in  those  from  Zaire.  The  numbers  of 
specimens  available  are  too  low  for  it  to  be  certain  that  these  distinctions  are  not  artefacts  of  sampling. 

DISTRIBUTION.  Records  suggest  that  the  species  occurs  at  high  altitudes  in  three  separate  areas:  Ethiopia; 
near  Nairobi  in  Kenya;  and  on  the  common  borders  of  Zaire,  Uganda,  Tanzania  and  Rwanda. 

MATERIAL  EXAMINED 

Zaire:  1  cf  (lectotype),  Ituri,  Nioka,  7.vii.l934  (Leroy)  (labelled:  'holotypus')  (MRAC). 

Ethiopia:  1  cf ,  Mt  Chilalo,  forest,  ca  8500  ft  (2550  m),  24.xi.1926  (Scott);  1  cf ,  Gojjam,  Fasilo  Bahr  Dar, 
scrub,  2.viii.l965  (Kitching);  1  cf,  Addis  Ababa,  Filoa,  vii.1941  (Meneghetti);  1  $,  Simien,  Atgheba 
Ghiyorghis,  ca  10,900  ft  (3270  m),  4.xii.l952,  arable  land  (Scott);  1  ? ,  Urgessa  R.,  Jimu,  2300  ft  (690  m), 
3.V.1905  (Zaphiro);  1  $,  Mt  Zuqala,  in  crater,  ca  9000  ft  (2700  m),  26.X.1926  (Omer  Cooper)  (BMNH). 
Zaire:  3  $  (paralectotypes),  Ituri,  Nioka,  vii.1934  (Leroy);  1  $  (paralectotype),  Ituri,  Nioka,  7.vii.l934 
(Leroy);  1  cf ,  Nioka,  vii.1934  (Leroy);  1  cf ,  Tshamgussa,  8-15.  viii.  1934  (de  Witte);  1  $,  Pare  Nat.  Albert, 
Lac  Gando,  2600  m,  Mt  Tamira,  ll.iii.1935  (de  Witte);  1  cf ,  Kivu,  Mulungu-Tshibinda,  xi.1951  (Lefevre) 
(MRAC);  1  $,  Ituri  Forest,  40  miles  (64  km)  NNE.  of  Beni,  3000  ft  (900  m),  10-12.ix.1959,  indigenous 
forest  with  cultivated  patches  (Cambridge  Expedition);  1  $ ,  Butembo-Beni  road,  29°30'E,  0°25'N,  4000  ft 
(1200  m),  ll.ix.1959,  agricultural  district  (Cambridge  Expedition)  (BMNH).  Uganda:  1  cf,  Kasinga 
Channel,  Katunguru,  1931  (Worthingtori)  (BMNH).  Kenya:  1  cf ,  Limuru,  10.iii.1911  (Anderson);  1  cf , 
Kinangop,  iii.1930  (Turner);  2  cf ,  W.  Aberdares,  above  9000  ft  (2700  m),  iii-iv.1934  (Turner);  1  $,  SE. 
slopes  of  Mt  Kenya,  6000-7000  ft  (1800-2100  m),  3-12.ii.1911  (Neave)  (BMNH).  Rwanda:  1  cf, 
Rubengera,  1900  m,  terr.  Kibuye,  12.ii.1953  (Basilewsky)  (MRAC).  Tanzania:  1  $,  Mt  Mem,  Olkokola, 
facing  NW. ,  2800  m,  24.vi-l.viu.1957  (Basilewsky,  Leleup)  (MRAC).  No  data:  1  cf  (ex  coll.  Schouteden) 
(MRAC). 


202  W.  R.  DOLLING 

Myla  microphthalma  Linnavuori 

(Figs  82, 105, 115, 117) 

Myla  microphthalma  Linnavuori,  1971:  175, 176-177.  Holotype  cf ,  IVORY  COAST:  Lamto,  3.iii-21.iv.l967 
(Gillon)  (R.  Linnavuori  coll.)  [not  examined]. 

Length:  cf,  8-3-9-7  mm;  $,8-6-9-9 mm. 

Head  (Fig.  82)  more  elongated  and  more  acutely  pointed  anteriorly  than  is  usual  in  the  genus,  eyes  small. 
Length  of  antennal  segment  I  divided  by  width  of  head  including  eyes  in  males  from  the  type-locality  (Ivory 
Coast)  0-91-0-97,  in  females  from  type-locality  0-90-0-97,  in  males  and  females  from  elsewhere  respective- 
ly 0-76-0-93  and  0-76-0-91.  Ratio  of  lengths  of  antennal  segments  in  males  and  females  from  Ivory  Coast 
respectively  about  l-00:0-86:l-05:0-98  and  l-00:0-88:l-04:0-85;  in  males  and  females  from  elsewhere 
respectively  about  1-00:0-88:1-05:1-17  and  1-00:0-90:1-05:1-03.  Length  of  rostral  segment  I  divided  by 
width  of  head  including  eyes  in  both  sexes  0-72-0-84;  ratio  of  lengths  of  rostral  segments  about 
l-00:0-85:0-44:0-67. 

Pronotum  (Fig.  82)  with  posterolateral  angles  rather  strongly  produced  anterolaterally  and  lateral 
margins  finely  granulate  with  a  few  larger  granules;  width  of  pronotum  across  apices  of  posterolateral 
spines  divided  by  width  of  head  including  eyes  2-15-2-65,  not  differing  significantly  between  the  sexes. 
Length  of  posterior  tibia  divided  by  length  of  posterior  femur  0-89-0-98. 

Abdomen  with  posterolateral  angles  of  sternites  III  to  VII  produced  as  short,  acute  spines.  Paramere 
(Fig.  105)  with  blade  broad,  distal  margin  of  apical  tooth  evenly  curved,  apical  tooth  rather  longer  than 
basal  tooth.  Spermatheca  (Fig.  115)  with  bulb  short,  duct  tightly  convoluted.  Second  valvula  of  ovipositor 
(Fig.  117)  apically  narrow,  elongate,  parallel-sided. 

Coloration  reddish  or  greyish  brown,  dorsal  midline  of  head  always  with  median  fuscous  stripe; 
pronotum  with  anterolateral  margins,  posterolateral  spines  and  anterior  half  of  midline  fuscous.  Colour 
pattern  otherwise  and  pubescence  as  in  M.  hoploxys. 

REMARKS.  The  fuscous  midline  of  the  head  distinguishes  this  species  from  all  others  in  the  genus.  The  series 
from  the  type-locality  (Ivory  Coast:  Lamto,  Toumodi)  was  found  to  differ  from  the  rest  of  the  material 
examined  in  several  biometric  ratios,  as  indicated  above.  A  particularly  striking  difference  was  found  in  the 
ratio  of  the  lengths  of  antennal  segments  I  and  IV.  The  length  of  segment  IV  divided  by  that  of  segment  I  in 
five  males  and  two  females  from  the  Ivory  Coast  was,  respectively,  0-91-1-00  and  0-84-0-86;  in  12  males 
and  six  females  from  the  rest  of  Africa  this  value  was,  respectively,  1-02-1-30  and  1-00-1-10.  Although  the 
Ivory  Coast  sample  size  is  small,  the  measurements  of  this  ratio  fall  completely  outside  the  range 
determined  for  specimens  from  the  rest  of  Africa.  It  may  be  that  the  Ivory  Coast  population  has  been 
isolated  from  the  remainder  for  a  considerable  period.  Perhaps  an  ancestral  population  of  this  species  was 
isolated  in  mountain  refugia  in  east  Central  Africa  and  Guinea  and  has  re-invaded  the  lower-lying  areas 
from  these  two  centres.  One  male  is  omitted  from  the  description  above  because  of  its  even  more  elongate 
head  and  its  aberrant  measurements.  It  is  from  Zaire:  Mongende;  its  length  is  9-0  mm;  the  width  of  its 
pronotum,  length  of  antennal  segment  I  and  length  of  rostral  segment  I,  all  divided  by  the  width  of  the 
head,  are  respectively  2-82,  1-03  and  0-84.  The  lengths  of  the  four  antennal  segments  are  in  the  ratio  of 
1-00:0-84:0-97:1-00. 

DISTRIBUTION.  The  species  is  known  from  two  widely  separated  areas:  (1)  Ivory  Coast  and  (2)  southern 
Africa,  avoiding  desert  and  dense  forest  and  extending  northwards  to  4°S  in  the  west  and  2°N  in  the 
equatorial  mountains  of  the  east. 

MATERIAL  EXAMINED 

Ivory  Coast:  6  cf,  3  $,  Lamto  (Toumodi),  5.vi.l962,  12.vi.1962,  10.vii.1962,  4.vi.l963,  ll.vi.1963, 
27.viii.1963,  21.vii.1964  (Gillon);  1  $,  Bouake,  21.vii.1962  (Gillon)  (BMNH).  Kenya:  1  cf,  Kaimosi, 
iii-iv.1932  (Turner)  (BMNH).  Rwanda:  1  cf  (paratype),  Kibungu,  1500  m,  2.ii.l953  (Basilewsky) 
(MRAC).  Urundi:  1  cf  (?paratype  -  locality  but  not  date  cited  in  original  description),  Kitega,  xi.1935 
(Lefevre);  1  $  (paratype),  Kitega,  vii.1935  (Lefevre);  2  $  (paratypes),  Usumbura,  1934  (Lefevre) 
(MRAC).  Zaire:  1  $,  Uele,  Wango,  29.vii.1931  (Bredo);  1  $,  Ituri,  Nioka,  vii.1934  (Leroy);  1  cf ,  Kivu, 
Mulungu,  1938  (Hendrickx);  1  cf,  Mayidi,  1942  (van  Eyen);  1  cf,  Kisantu,  1919  (Vanderijst)  (all 
paratypes);  1  cf ,  Tshambi,  xii.1934  (de  Witte);  1  $,  Reg.  Thysville  (Bas-Congo),  1959-1963  (Michaux);  1 
$,  Lulua,  Kapanga,  i.1933  (Overlaet);  1  cf,  Katanga,  Kipopo,  29.X.1961  (Marechal)  (MRAC);  2  $>, 
Elisabethville  (=  Lubumbashi),  25.iv.1939,  15.xi.1939  (Bredo)  (IRSNB).  Zambia:  1  cf,  Kamankundju 
River,  Ikelenge,  N.  Mwinilunga,  5.V.1972  (Pinhey,  de  Moor)  (NMB).  Angola:  1  cf ,  12  miles  (19  km)  SW. 
of  Luimbale,  ca  1680  m,  20-2 l.iii.  1972,  general  sweeping  (BMNH  Southern  African  Expedition) 
(BMNH).  Malawi:  1  cf ,  SE.  shore  Lake  Nyasa,  between  Fort  Maguire  and  Fort  Johnston,  6-17. iii.  1910 


TRIBE  PSEUDOPHLOEINI 


203 


111 


112 


114 


118 


115 


Figs  111-121  Myla  species.  111-114,  calida:  (111)  dorsal  view  of  phallotheca,  conjunctiva  and  vesica; 
(112)  lateral  view  of  same,  dorsal  to  left;  (113)  ventral  view  of  same;  (114)  apical  view  of  same.  115, 116, 
spermatheca  of  (115)  microphthalma;  (116)  onceroma.  117-121,  left  lateral  view  of  apex  of  left  second 
valvula  of  ovipositor  of  (117)  microphthalma;  (118)  onceroma;  (119)  hoploxys;  (120)  calida;  (121) 
granula. 


(Neave)  (BMNH).  South  Africa:  1  cf ,  Transvaal,  Pretoria,  26.xi.1973  (Jacobs);  1  cf ,  Utrecht,  4.xi.l967 
(Kroon);  1  $,  Camperdown,  17.iv.1908  (Leigh)  (TM);  1  cf ,  Transvaal,  10 miles  (16 km)  NE.  of  Cullinan, 
27.ii.1962  (Capener);  1  <j>,  no  precise  data  (Capener)  (J.  A.  Slater  coll.);  1  cf,  Umbilo  (Bevis);  2  $, 
Transvaal,  Argent,  29.iii.1942  (Capener);  1  $,  Pondoland,  Port  St  John,  15-31.V.1923  (Turner);  1  cf, 
Natal,  Tugela  River,  nrWeenen,iii.  1897;  1  cf,  Natal,  Karkloof,  10.ii.l897(?or2.x.l897);  1  cf,  1  $, Natal, 
Howick;  1  cf ,  Natal,  Estcourt;  3  cf ,  2  $,  Estcourt,  xi.1896;  4  cf ,  Estcourt  (Marshall)  (BMNH).  Possibly 
belonging  to  this  species:  Zaire:  1  cf ,  Mongende,  17. iv. 1921  (Schouteden)  (falsely  labelled  as  a  paratype  of 
Myla  gracilis  Schouteden  but  not  from  a  locality  mentioned  in  the  original  description)  (MR AC). 

Myla  onceroma  sp.  n. 

(Figs  80, 116, 118) 

Length:  cf ,  8-2-9-2 mm;  $,8-6 mm. 

Head  (Fig.  80)  not  elongate  or  acute  anteriorly.  Eyes  small.  Length  of  antenhal  segment  I  divided  by 
width  of  head  including  eyes  in  male  0  •  73-0  •  84 ,  in  female  (one  only  seen)  0  •  78 .  Ratio  of  lengths  of  antennal 
segments  in  male  about  1-00:0-83:1-02:1-19,  in  female  as  1-00:0-81:1-04:1-04.  Length  of  rostral  segment  I 
divided  by  width  of  head  including  eyes  in  male  0-70-0-75,  in  female  0-78.  Ratio  of  lengths  of  rostral 
segments  in  male  about  l-00:0-82:0-43:0-65,  in  female  as  l-00:0-77:0-42:0-63. 

Pronotum  (Fig.  80)  with  posterolateral  angles  shortly  produced  anteriorly,  bearing  short  spines;  width 
across  apices  of  spines  divided  by  width  of  head  including  eyes  in  male  1-99-2-20,  in  female  2-03.  Length  of 
posterior  tibia  divided  by  length  of  posterior  femur  0-94-0-97. 

Abdominal  sternites  III  to  VII  with  posterolateral  angles  produced  into  short,  acute  spines.  Male 
genitalia  similar  to  those  of  M.  microphthalma.  Female  with  bulb  of  spermatheca  (Fig.  116)  longer  than 
that  of  M.  microphthalma  and  with  apical  projection  of  second  valvula  (Fig.  118)  very  short. 

General  coloration  dark  red-brown  or  brown.  Midline  of  head  and  pronotum  concolorous,  without 
darker  median  stripe. 

REMARKS.  This  small  species  is  probably  most  closely  related  to  M.  microphthalma  but  may  be  disting- 
uished from  it  by  the  much  shorter  antennae,  shorter  head  and  much  less  strongly  produced  pronotal 
angles,  which  approach  the  form  of  those  of  M.  niokensis  but  bear  longer  apical  spines.  It  is  a  montane 


204  W.  R.  DOLLING 

form,  perhaps  descended  from  the  ancestral  population  that  gave  rise  to  M.  microphthalma  but  is  now 
greatly  specialized,  particularly  in  the  form  of  the  second  valvulae  of  the  ovipositor,  after  long  isolation. 

DISTRIBUTION.  Known  only  from  three  localities  at  high  altitudes  in  Cameroun  and  south-eastern  Nigeria. 

MATERIAL  EXAMINED 

Holotype  cf ,  Cameroun:  Bamenda,  17.xii.1955  (Bechyne,  Exped.  Mus.  G.  Frey}  (BMNH). 

Paratypes.  Cameroun:  4  cf,  as  holotype  but  dates  7.xii.l955,  8.xii.l955,  13.xii.1955  and  17.xii.1955 
(BMNH);  2  Cf ,  1  $ ,  Mt  Bamboutos  (MRAC). 

Non-paratypic  material  (received  after  description  was  drawn  up).  Nigeria:  3  d",  4  9 ,  SE.  State,  Obudu 
C.R.,  21  and  23.iii.1971  (Medler)  (University  of  Ibadan). 

Myla  gracilis  Schouteden 
(Figs  83,  94,  98, 108-110) 

Myla  gracilis  Schouteden,  1938:  293.  LECTOTYPE  cf ,  ZAIRE  (MRAC),  here  designated  [examined]. 
Myla  graciloides  Linnavuori,   1971:   175.  Lectotype  cf,  SUDAN:  Equatoria  prov.,  Yei-Maridi,  13- 

15. iv. 1963,  swampy  meadow  (Linnavuori)  (R.  Linnavuori  coll.),  designated  by  Linnavuori,  1978:  36 

[not  examined].  Syn.  n. 

Length:  cf,  8-4-9-8  mm;  $,  9-00-10-0  mm. 

Head  distinctly  longer  than  wide,  narrowly  rounded  anteriorly.  Length  of  antennal  segment  I  divided  by 
width  of  head  including  eyes  in  male  1-06-1-27,  in  female  1-06-1-25.  Ratio  of  lengths  of  antennal  segments 
in  male  about  l-00:0-87:0-94:0-86,  in  female  about  1-00:0-87:0-93:0-80.  Length  of  rostral  segment  I  divided 
by  width  of  head  including  eyes  in  both  sexes  0-71-0-82.  Ratio  of  lengths  of  rostral  segments  about 
l-00:0-85:0-46:0-66. 

Pronotum  (Fig.  83)  with  posterolateral  angles  strongly  produced  anterolaterally,  terminating  in  slender 
spines,  anterolateral  margins  with  one  to  three  prominent,  piceous  or  black  tubercles;  width  across  apices 
of  posterolateral  spines  divided  by  width  of  head  including  eyes  in  male  2-57-3-18,  in  female  2-74-3-09. 

Abdomen  with  posterolateral  angles  of  sternites  III  to  VII  produced  as  fine  spines  (Fig.  94).  Connex- 
ivum  rather  narrow;  laterotergite  V  twice  as  long  as  wide.  Paramere  (Figs  108-110)  slender,  with  apical 
tooth  long  and  both  teeth  on  a  common  stem  that  arises  terminally  or  subterminally  from  the  blade.  Female 
with  second  valvula  tapering  to  narrowly  rounded  apex. 

Pubescence  of  head,  pronotum,  scutellum,  thoracic  pleura,  abdominal  venter  laterally  and  basally, 
laterotergites,  clavus,  corium,  antennal  segment  I,  coxae,  trochanters,  femora  and  rostrum  comprising 
fairly  short,  flattened,  scale-like,  decumbent  (except  on  appendages),  white  hairs;  hairs  on  thoracic  pleura 
(Fig.  98)  and  declivent  area  of  pronotum  very  distinctly  longer  than  distance  between  adjacent  hair 
insertions  and  tending  to  form  three  or  four  transverse  bands  on  deolivent  area  of  pronotum.  Areas  of 
abdominal  venter  and  appendages  that  are  devoid  of  scale-like  pubescence  with  short,  suberect,  colourless 
hairs. 

Coloration  generally  as  in  M.  hoploxys  but  never  heavily  tinged  red.  Scutellum  with  midline  and  small, 
apical  knob  usually  creamy  white.  Large  tubercles  of  anterolateral  margins  of  pronotum  and  apices  of 
posterolateral  spines  piceous  to  black.  Veins  of  hemelytral  membrane  brown,  uninterrupted. 

REMARKS.  This  species  differs  from  all  others  of  the  genus  in  the  greater  length  of  the  scale-like  pubescence. 
The  posterolateral  pronotal  angles  are  more  strongly  produced  in  this  species  than  in  any  of  its  congeners. 
The  combination  of  these  two  features  gives  M.  gracilis  a  superficial  resemblance  to  Clavigralla  schnelli 
(Villiers).  Indeed,  the  type-series  of  M.  gracilis  contains  three  individuals  of  C.  schnelli,  one  of  which  bears 
a  label  with  the  legend,  'holotypus'.  This  type  designation  has  never  been  published  and  Schouteden's  very 
detailed  description  of  the  antennifers,  rostrum,  antennae,  femoral  spines  and  male  genital  capsule  leaves 
no  doubt  that  he  was  describing  a  species  of  Myla.  I  have,  therefore,  ignored  the  unpublished  'holotype' 
designation  and  here  select  one  of  the  'paratypes'  as  lectotype.  Linnavuori  (1971:  175;  1978:  35-36)  used 
the  name  M.  gracilis  for  what  is  evidently  C.  schnelli  and  established  the  unnecessary  new  nominal  species 
Myla  graciloides  for  M.  gracilis  proper.  In  his  1971  paper,  Linnavuori  cites  the  localities  Sudan,  Congo  and 
Ivory  Coast  but  in  1978  he  cites  only  the  'type'  from  Sudan;  this  citation  is  effectively  a  lectotype 
designation.  There  is  some  variation  in  the  pronotal  posterolateral  spines  which  are  usually  laterally 
directed  but  sometimes  anterolaterally  directed. 

DISTRIBUTION.  West  and  Central  Africa. 


TRIBE  PSEUDOPHLOEINI 


205 


MATERIAL  EXAMINED 

Zaire:  1  cf  (lectotype  of  gracilis),  Yangambi,  xii.1925  (Ghesquiere)  (labelled  as  paratype  on  pink 
rectangle  but  lectotype  on  purple-bordered  disc)  (MRAC) 

Ivory  Coast:  14  cf,  20  $,  Lamto,  variously  'Lamto  (Toumodi)',  various  dates  from  4.xii.l962  to 
14.viii.1964,  all  months  except  i,  ii,  xi,  greatest  numbers  in  vi;  5  cf ,  1  $ ,  Bouake,  22.V.1962, 21.vii.1962;  1 
CT,  Dabov,  S.v.1964  (BMNH).  Ghana:  1  cf,  'Gold  Coast'  (CottereU);  1  $,  Yapi  (Simpson};  1  $,  Tafo, 
9.V.1957,  in  yellow  trays  (Eastop)  (BMNH);  2  cT,  1  ? ,  Tafo,  14.X.1966, 20.ii.1967,  pyrethrum  knockdown 
from  cocoa  (Kumar,  Leston);  1  $ ,  Legon,  2.x.  1968  (Kumar);  1  $ ,  Ho,  7.xi.l967  (Leston);  1  $ ,  Shai  Hill, 
29.vii.1967  (Leston)  (UG).  Togo:  1  cf,  Missahou<§,  650  m,  vi.1963  (Schach)  (MRAC).  Nigeria:  1  $, 
Akpashe  District,  Udi,  27.X.1955  (Bechyne)  (BMNH).  Zaire:  73  cf,  61  $,  Faradje,  29°40'E,  3°40'N, 
i.1913, 3  $, Faradje, i.1912;  1  cf ,  1  $, Faradje, i.1917;  1  cf , Medje, 27°15'E, 2°25'N,  10.iv.1910 (all Lang, 
Chapin)  (AMNH);  1  cf,  Lomani,  xii.1931  (Collart);  1  cf,  Ngowa,  vi.1938  (Mertens);  1  $,  Ngowa, 
7-27.iv.1939  (Mertens)  (IRSNB);  2  $,  Congo  da  Lemba,  1911  (Mayne);  1  $,  Haut-Uele,  Moto,  1920 
(Burgeon);  1  $ , Elisabethville  (R.  Lubumbashi),  7.xi.l920 (Bequaert);  1  $,  Katanga,  Kando,  l-4.iv.1931 
(de  Witte);  1  $,  Kisanfu,  1919  (Vanderijst);  1  ex.  without  abdomen,  Magabi-Niarembe,  v.1935  (Scops) 
(these  seven  exx  all  paralectotypes  of  gracilis);  1  cf ,  Kivu,  Bwito,  ca  1700  m,  26. vi. 1934  (Marlier);  1  cf, 
Ituri,  Bunia,  vi.1938  (Lefevre);  1  cf,  Bas-Congo,  Mayidi,  1942  (van  Eyen);  1  cf,  Rutshuru,  Fuku, 
30.V.1936  (Lippens);  1  cf ,  Lulua,  Kapanga,  Hi. 1933  (Overlaet)  (MRAC). 

Paralectotypes  of  My  la  gracilis  Schouteden  not  belonging  to  this  species.  3  exx  determined  as  Clavigralla 
schnelli  (Villiers)  by  Dolling,  1979:  36,  q.v. 

Distribution  of  Pseudophloeinae 

Preamble 

Without  a  complete  phylogenetic  analysis  of  the  subfamily,  incorporating  the  Palaearctic 
genera,  a  comprehensive  account  of  its  zoogeography  is  not  possible.  In  undertaking  the  present 
revision  I  have  considered  every  species  of  the  subfamily,  including  the  Palaearctic  ones,  in 
relation  to  the  diagnostic  features  of  every  genus,  and  I  am  reasonably  certain  that  all  of  the 
genera  currently  recognized  are  monophyletic  in  origin  and  holophyletic  in  composition. 
However,  I  have  not  fully  investigated  the  relationships  among  all  the  genera  of  the  subfamily. 
Two  presumably  holophyletic  groups  of  genera  are  recognized:  the  tribe  Clavigrallini  (see 
Dolling,  1978;  1979a)  and  a  large  group  of  genera  including  all  of  those  confined  to  the  North 
Temperate  Zone  and  all  of  the  tropical  genera  except  for  Vilga,  Risbecocoris,  Paramyla, 
Hoplolomia  and  Indolomia.  The  first  two  of  these  genera  are  probably  not  descended  from  the 
most  recent  common  ancestor  of  the  two  big  groups  while  the  last  three  may  be  related  to  the 
non-clavigralline  group  in  some  way  (see  Table  1).  The  distribution  of  the  various  genera  of 
Pseudophloeinae  in  the  major  zoogeographical  regions  is  summarized  in  Table  2.  The  Afro- 
tropical  region  in  this  table  excludes  the  Mascarene  area  (Madagascar,  Mauritius,  Reunion  and 
Rodriguez),  which  is  shown  separately.  The  Oriental  region  is  taken  to  include  New  Guinea  and 
the  Solomon  Islands  but  Australia  is  excluded  and  its  pseudophloeine  fauna  is  tabulated  in  a 
separate  column. 

Table  1     Distribution  of  two  characters  in  the  subfamily  Pseudophloeinae. 


Antevannal  vein 
absent 


Antevannal  vein 
present 


Posterior  femur  without  basal  tubercle 
Posterior  femur  with  basal  tubercle 


Vilga, 
Risbecocoris 

Hoplolomia, 

Indolomia, 

Paramyla 


Clavigrallini 

Remaining 
genera 


Neotropical 

The  only  genus  of  Pseudophloeinae  represented  in  the  Neotropical  region  is  Vilga,  which  is 
endemic  to  the  New  World.  The  14  species  were  divided  among  six  subgenera  by  Dolling  (1977), 


206  W.  R.  DOLLING 

who  commented  that  the  differences  among  these  subgenera  were  of  the  same  degree  as  those 
separating  some  Palaearctic  genera.  One  subgenus,  whose  single  species  is  facultatively 
brachypterous,  occurs  from  the  extreme  southern  U.S.A.  to  El  Salvador;  all  the  other 
subgenera  are  confined  to  the  Neotropical  region  and  are  widely  sympatric  there,  with  the 
exception  of  one  subgenus  whose  three  species  are  confined  to  the  temperate  and  largely 
mountainous  regions  of  Argentina,  Chile  and  Peru.  This  latter  subgenus,  through  a  general 
reduction  of  spininess  of  the  body  and  legs  and  a  flattening  and  broadening  of  the  body, 
superficially  resembles  the  tropical  montane  and  temperate  South  African  genus  Paramyla  and 
the  Palaearctic  Arenocoris  and  Bathysolen  Fieber. 

Nearctic 

The  presence  of  a  single  species  of  the  otherwise  Neotropical  genus  Vilga  in  Mexico  and  adjacent 
areas  has  been  mentioned  above.  In  addition,  two  genera  of  Pseudophloeini  are  shared  between 
the  Palaearctic  and  Nearctic  regions. 

Coriomeris  Westwood  has  16  species  in  the  Palaearctic  and  another  four  in  the  Nearctic.  The 
Nearctic  species  (see  Dolling  &  Yonke,  1976)  are  mutually  very  similar  and  have  strictly 
parapatric  distributions,  suggesting  recent  entry  into  and  radiation  within  the  region.  Species  of 
Coriomeris  are  found  within  the  Arctic  Circle  in  Alaska,  Yakutia  and  Karelia,  so  there  would 
seem  to  be  no  climatic  barrier  to  migration  across  the  Bering  Straits  even  today. 

Ceraleptus  Costa  has  five  Palaearctic  and  four  Nearctic  species,  none  of  them  shared  between 
the  two  regions.  Its  nearest  relatives  appear  to  be  Microtelocerus  Reuter  and  Urartucoris 
Puchkov,  both  genera  with  very  restricted  distributions  in  the  mountains  of  the  central  southern 
Palaearctic,  so  a  Palaearctic  origin  for  Ceraleptus  seems  probable.  Its  American  species  are 
rather  diverse  and  fall  geographically  into  two  pairs,  one  pair  sympatric  along  the  Pacific  coast  of 
North  America  and  the  other  pair  sympatric  in  the  south-eastern  U.S.A.  (Froeschner,  1963). 
Members  of  the  genus  do  not  occur  at  such  high  latitudes  as  the  extremes  penetrated  by 
Coriomeris  and  this  fact,  coupled  with  the  greater  diversity  and  more  complex  distribution 
pattern  shown  by  the  species  of  Ceraleptus,  suggests  that  the  genus  has  been  present  in  the 
Nearctic  region  longer  than  has  Coriomeris,  presumably  arriving  in  one  or  more  immigration 
episodes  during  an  era  of  warmer  climate  than  that  of  the  present  day. 

Palaearctic 

Twelve  genera  and  about  39  species  of  Pseudophloeini  occur  in  the  Palaearctic  region;  only  one 
of  these  species  is  shared  with  another  region:  Arenocoris  intermedius ,  whose  range  includes 
western  temperate  Asia,  Iberia  and  the  northern  coastal  countries  of  Africa  and  extends 
southwards  into  Chad  and  Ethiopia.  As  indicated  above,  the  Palaearctic  genera  Coriomeris  and 
Ceraleptus  have  both  entered  North  America  and  have  endemic  species  there. 

The  Palaearctic  intrusions  of  two  species  of  the  tribe  Clavigrallini,  Clavigralla  scutellaris  (into 
Afghanistan)  and  Gralliclava  horrens  (into  Japan)  are  mentioned  below;  no  other  genus  of  the 
subfamily  is  represented  in  both  the  Palaearctic  and  Afrotropical  or  the  Palaearctic  and  Oriental 
regions. 

Afrotropical  and  Oriental  Clavigrallini 

The  well-defined  tribe  Clavigrallini  comprises  four  genera  whose  interrelationships  are  unclear. 
With  the  exception  of  two  minor  intrusions  into  the  Palaearctic  region,  mentioned  below,  the 
tribe  is  confined  to  the  tropical  regions  of  the  Old  World. 

Clavigralla  Spinola  is  the  largest  genus  of  Pseudophloeinae,  with  46  described  species.  It  has 
clearly  undergone  most  of  its  radiation  on  the  African  mainland,  where  38  of  its  species  are 
found.  Only  one  species,  C.  elongata  Signoret,  which  is  widespread  on  the  African  mainland,  is 
present  on  the  smaller  offshore  Afrotropical  islands.  It  has  reached  Sao  Tome  and  the  Cape 
Verde  Islands  in  the  Atlantic  (Dolling,  1979a:  28  erroneously  reported  it  from  the  Canary 
Islands  on  the  basis  of  a  mislabelled  specimen  from  the  Wollaston  collection;  the  card  bears  a 


TRIBE  PSEUDOPHLOEINI  207 

Table  2  Number  of  species  of  each  genus  of  Pseudophloeinae  in  the  regions  indicated.  Species  occurring 
in  more  than  one  region  are  scored  in  parentheses  in  regions  other  than  those  of  their  putative  origin  as 
are  genera  with  no  endemic  species. 


13 

u 

13 
u                  <u 

o 

e, 

.a           '§ 

'g.         § 

o 

13 

.2 

o 
j^ 

t"i                                                ^ 

5              c3 

13 

tn                                ^ 

2              8 

S 

"55                5 

<u 

QJ                                    ^ 

<i3 

•c 

2 

t   _                                       TO 

£                         OH 

<             S 

0 

<              H 

V%a                        13+(1) 

1 

14 

Ceraleptus 

4                5 

9 

Coriomeris 

4               16 

20 

Arenocoris 

4 

(1) 

4 

(nine  genera1) 

15 

15 

Risbecocoris 

3 

2 

5 

Hoplolomia 

2 

2 

Indolomia 

1 

1 

Pungra 

1 

1 

Pseudomyla 

2 

2 

Psilolomia 

4 

9 

13 

Paramyla 

2 

2 

Neomevaniomorpha 

1 

1 

Mevaniomorpha 

2 

2 

Mevanidea 

1               1 

2 

Myla 

11 

11 

Clavigralla 

(1) 

38           6+(l) 

2+(l) 

46 

Oncaspidia 

1 

1 

Clavigralloides 

5 

(2)               5 

Grallidava 

(1) 

8 

2               10 

Total  genera                  1 

3           12+  (2) 

9+(l)           2 

9 

1+(1)           28 

Total  species                 13 

9           40+  (2) 

63+(l)       7+(l) 

32+(l) 

2+  (2)          166 

1  Anoplocerus,  Bathysolen,  Bothrostethus,  Loxocnemis,  Microtelocerus,  Nemocoris,  Strobilotoma,  Ulmicola,  Urar- 
tucoris. 

colour-coded  mark  that  indicates  its  true  provenance  as  the  Cape  Verde  Islands),  and  Madagas- 
car, Mauritius,  Reunion,  Rodriguez  and  the  Seychelles  in  the  Indian  Ocean.  It  has  also  been 
reported  from  Arabia.  In  1979 1  had  seen  no  specimens  of  the  species  from  the  mainland  of  West 
Africa,  where  it  seems  to  be  largely  replaced  by  C.  shadabi  Dolling  and  C.  breviceps  Dolling.  I 
have  now  seen  specimens  of  it  from  two  localities  in  Senegal.  C.  elongata  is  the  only 
representative  of  the  subfamily  in  these  Atlantic  and  Indian  Ocean  islands  with  the  exception  of 
six  endemic  species  in  Madagascar.  One  of  these,  C.  tuberculicollis  (Reuter),  is  a  taxonomically 
isolated  species  with  several  plesiomorphic  features;  it  has  probably  been  present  in  Madagascar 
much  longer  than  any  other  species  of  the  tribe.  Three  Malagasy  species  are  closely  similar  to  C. 
elongata  and  may  be  either  local  derivatives  of  it  or  descendants  of  a  single,  elongata-like 
immigrant.  The  other  two  endemics  are  members  of  the  C.  tomentosicollis  Stal  group  but  belong 
to  different  subgroups  and  clearly  derive  from  two  separate  immigrations.  Thus,  Madagascar 
seems  to  have  been  colonized  on  at  least  four  and  possibly  five  separate  occasions  by 
Clavigrallini,  but  there  is  evidence  for  local  radiation  stemming  from  only  one  of  these  invasions. 
Clavigralla  is  evidently  a  recent  arrival  in  the  Oriental  region.  Its  three  Oriental  species  all 
belong  to  the  same  subgroup  of  the  tomentosicollis-group.  One  of  them,  C.  scutellaris  (West- 
wood),  which  is  probably  the  sister-species  of  C.  tomentosicollis,  is  also  found  in  north-eastern 
Africa  and  Arabia;  in  Asia  it  extends  as  far  northwards  as  Afghanistan.  The  other  two  are 
vicariant  sister-species,  one  confined  to  Sri  Lanka  and  southern  India  and  the  other  distributed 
from  northern  India  to  southern  China  and  Vietnam;  the  Indian  population  of  this  second 
species  is  recognizably  distinct  from  the  populations  further  east. 


208  W.  R.  DOLLING 

Oncaspidia  Stal  has  a  single,  widespread  species  endemic  to  Africa. 

Clavigralloides  Dolling  has  the  most  easterly  centre  of  distribution  of  any  genus  of  Pseudo- 
phloeinae.  Three  of  its  five  species  occur  in  New  Guinea;  one  of  these  three  is,  so  far  as  is  known, 
endemic  while  the  other  two  extend  southwards  into  northern  Australia  and  one  of  these  also 
spreads  northwards  to  southern  China.  Closely  related  to  these  two  is  a  species  restricted  to  the 
Lesser  Sunda  Islands.  The  fifth  member  of  the  genus  is  rather  isolated  taxonomically  and  is 
divided  into  northern  Indian  and  a  southern  Chinese  subspecies. 

Gralliclava  Dolling  species  are  distributed  from  India  to  Australia.  They  fall  into  two  groups, 
one  with  a  plesiomorphic  condition  of  the  male  genital  capsule  and  a  variety  of  unusual 
paramere  shapes,  and  the  other  with  an  apomorphic  condition  of  the  capsule  and  rather  uniform 
parameres.  The  first  group  comprises  six  known  species  with  restricted  distributions;  three  of 
these  are  Indian;  one  is  found  in  the  Himalayas  and  at  high  altitudes  in  southern  China  and  Laos, 
one  is  known  only  from  its  high  altitude  type-locality  in  Java  and  one,  surprisingly,  is  known 
from  a  single  specimen  taken  on  a  small  Australian  island  in  the  Torres  Strait.  The  four  species  of 
the  second  group  between  them  occupy  an  almost  continuous  range  from  Sri  Lanka  and  India  to 
Japan,  the  Solomon  Islands  and  tropical  Australia.  One  is  known  from  two  specimens,  one  from 
Timor  and  the  other  from  Flores.  The  most  widespread  species,  G.  horrens  Dohrn,  is  also 
known  from  Timor,  from  Wetter  and  the  Sulawesi  group,  and  thence  northwards  to  Japan 
(Japanese  records,  all  from  Kyushu,  are  summarized  by  Kawazawa,  1978)  and  westwards  to 
India  and  Sri  Lanka.  A  third  species  is  common  in  New  Guinea  and  is  also  recorded  from 
Ambon  and  the  Admiralty,  Bismarck  and  Solomon  groups.  The  fourth  species  has  been 
collected  from  many  localities  in  Queensland  and  the  Northern  Territory  of  Australia  but  it  has 
never  been  found  elsewhere,  nor  has  any  other  species  of  the  genus  been  taken  in  Australia.  The 
almost  complete  allopatry  of  these  four  species  and  the  fact  that  the  Torres  Strait  separates  two 
of  them  suggests  a  very  recent  radiation  of  this  group. 

Afrotropical  and  Oriental  Pseudophloeini 

Apart  from  the  intrusive  Arenocoris  intermedius ,  the  present  revision  recognizes  seven  genera 
with  25  species  in  the  Afrotropical  region  and  six  genera  with  17  species  in  the  Oriental.  Two  of 
these  genera  and  none  of  the  species  are  shared  between  the  two  regions.  The  largest  genera  are 
the  wholly  Afrotropical  My  la,  with  11  species,  and  the  shared  Psilolomia,  with  13. 

Risbecocoris ,  with  five  described  species,  occupies  a  probably  almost  continuous  belt  of 
grassland  and  semi-desert  from  Senegal  via  Arabia  to  Delhi,  taking  in  the  Air  mountains  and 
northern  Kenya  as  well.  This  type  of  habitat  has  probably  been  available  in  the  area  for  a  very 
long  time  and  dispersal  across  it  between  Africa  and  Asia  would  present  no  problems  for  an 
organism  as  well  adapted  to  it  as  is  Risbecocoris.  It  makes  little  sense  to  attribute  part  of  this  arid 
zone  to  the  Afrotropical  region  and  part  to  the  Oriental  in  the  context  of  the  biogeography  of 
such  organisms. 

Psilolomia  displays  its  greatest  diversity  in  the  Oriental  region.  A  group  of  dark-coloured 
species,  whose  characteristic  habitat  is  probably  forest  clearings,  extends  no  further  westwards 
than  eastern  India  while  a  group  of  yellowish  species,  probably  characteristic  of  dry,  open 
woodland,  is  distributed  from  West  Africa  to  South  East  Asia.  Only  four  species  of  this  group 
occur  in  Africa;  they  have  had  little  or  no  success  south  of  the  Equator.  One  of  them,  Psilolomia 
nigeriensis,  shows  a  tendency  to  split  into  northern  and  southern  races  in  Nigeria,  perhaps  in 
response  to  differing  climatic  conditions.  A  single  migration  in  the  fairly  recent  past  from  India 
into  Africa  via  a  more  humid  Arabia  seems  the  most  probable  explanation  for  its  occurrence  in 
both  continents.  Large  areas  of  apparently  suitable  terrain  remain  unoccupied  by  this  genus  in 
eastern  and  southern  Africa. 

Mevanidea  is  the  only  genus  of  Pseudophloeinae,  apart  from  Clavigralla,  to  have  been  found 
in  Madagascar.  The  two  species  of  the  genus  show  classic  allopatry,  one  in  mainland  Africa  and 
the  other  endemic  to  Madagascar. 


TRIBE  PSEUDOPHLOEINI  209 

Discussion 

The  faunistic  picture  that  emerges  is  one  of  greatest  diversity  in  the  tropics  of  the  Old  World  with 
rather  less  diverse  but  long  established  faunas  in  the  Palaearctic  and  Neotropical  regions.  The 
Neotropical  and  Palaearctic  faunas  seem  to  have  developed  in  almost  complete  isolation  from 
each  other  and  from  the  Palaeotropical  faunas.  The  Nearctic  region,  Australia  and  Madagascar 
have  no  endemic  genera  and  have  played  only  a  peripheral  part  in  the  evolutionary  history  of  the 
subfamily.  Apart  from  Risbecocoris,  whose  range  is  in  a  transitional  zone,  only  one  genus, 
Psilolomia,  is  shared  between  the  Oriental  and  Afrotropical  regions.  It  is  the  only  genus  that  can 
be  shown  to  have  successfully  invaded  and  begun  to  diversify  in  a  region  where  a  diverse 
pseudophloeine  fauna  was  already  present. 

The  genus  Vilga  has  evidently  been  present  in  the  Neotropics  for  a  very  long  time,  perhaps 
since  before  the  opening  of  the  Atlantic  Ocean;  this  would  place  the  early  radiation  of  the 
subfamily  before  the  end  of  the  Cretaceous  period.  Among  tropical  Coreoidea  it  is  frequently 
the  case  that  the  distribution  of  the  subfamilies  is  pantropical  while  the  tribes  are  restricted  to 
either  the  Old  World  or  the  New. 

Dense  forest  and  water  seem  to  have  been  very  effective  barriers  to  dispersal  of  Pseudo- 
phloeini  but  less  effective  in  the  case  of  Clavigrallini.  The  more  elongate  condition  of  the 
hemelytra  of  the  latter,  with  a  tendency  for  the  corium  to  extend  apically  along  the  costal  margin 
of  the  hemelytron,  and  the  deeper  thorax,  presumably  housing  larger  flight-muscles,  are 
suggestive  of  a  more  sustained  power  of  flight  which  would  enable  its  possessor  to  overcome  the 
obstacles  that  inhibit  the  dispersal  of  Pseudophloeini.  The  distribution  patterns  of  two  groups  of 
recently  differentiated  species  illustrate  this  difference  in  dispersal  ability,  and  hence  in  rates  of 
genetic  interchange.  The  genus  Coriomeris  in  North  America  has  split  into  four  recognizably 
distinct  though  closely  related  species  without  any  intervening  barriers  of  forest  or  water, 
whereas  the  species  of  the  horrens-group  of  the  clavigralline  genus  Gralliclava  are,  with  one  very 
minor  exception,  separated  by  sea  barriers.  Parapatric  speciation  may  also  be  in  progress  in  the 
case  of  the  northern  and  southern  forms  of  Psilolomia  nigeriensis. 

Acknowledgements 

I  am  sincerely  grateful  to  Dr  Roberto  Poggi,  Museo  Civico  di  Storia  Naturale,  Genoa,  for  sending  me  a 
drawing  of  the  holotype  of  Pseudophloeus  gestroi;  to  Prof.  Dott.  Carlo  Vidano,  Turin,  for  searching 
Spinola's  collection  for  the  type-material  of  Merocoris  spinicollis;  to  Dr  H.  Brailovsky,  UNAM,  Mexico, 
for  donating  the  unique  type-specimen  of  Indolomia  conculata;  and  to  the  following  colleagues  for  their 
kindness  in  arranging  loans  of  material  in  the  depositories  mentioned:  Dr  A.  E.  Akingbohungbe,  Ife 
University,  Ile-Ife,  Nigeria;  Dr  P.  H.  Arnaud,  CAS,  San  Francisco;  Mr  J.  C.  Deeming,  formerly  IAR, 
Samaru;  Dr  U.  Gollner-Scheiding,  MNHU,  Berlin;  Mrs  C.  N.  Higa  and  the  late  Dr  J.  L.  Gressitt,  BPBM, 
Honolulu;  Dr  A.  Kaltenbach,  NMV,  Vienna;  Prof.  R.  Kumar,  UG,  Legon;  Drs  I.  Lansbury  and  M.  W.  R. 
de  V.  Graham,  UM,  Oxford;  Dr  R.  Linnavuori,  Raisio,  Finland  (private  collection  to  be  deposited  in 
ZMU,  Helsinki);  Dr  J.  G.  H.  Londt,  NMP,  Pietermaritzburg;  Dr  J.  T.  Medler,  University  of  Ibadan, 
Nigeria;  Drs  M.  Meinander  and  A.  Jansson,  ZMU,  Helsinki;  Dr  J.  Nast,  IZPAN,  Warsaw;  DrT.  Nyholm, 
NR,  Stockholm;  Dr  G.  Peterson,  IP,  Eberswalde;  Dr  E.  C.  G.  Pinhey,  NMB,  Bulawayo;  Mr  P.  Reavell, 
Empangeni,  Zululand  (private  collection,  type-material  to  be  deposited  in  NMP,  Pietermaritzburg);  Dr  G. 
Schmitz,  MRAC,  Tervuren  and  IRSNB,  Brussels;  Drs  M.  U.  Shadab  and  P.  Wygodzinsky,  AMNH,  New 
York;  Prof.  J.  A.  Slater,  University  of  Connecticut,  U.S.A.  (private  collection);  Dr  M.  Taou  and  the  late 
Dr  A.  Villiers,  MNHN,  Paris;  Mr  J.  A.  van  Reenen,  TM,  Pretoria. 

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Entomologique  de  Belgique  53:  250-309;  27  figs. 

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+  438  pp.  London. 

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figs. 
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the  British  Museum  (Natural  History)  (Entomology)  36:  281-321, 108  figs. 
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checklist  of  the  world  species.  Bulletin  of  the  British  Museum  (Natural  History)  (Entomology)  39: 1-84, 

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1979c.  A  second  species  of  Microtelocerus  Reuter  (Hem. ,  Coreidae)  with  a  redescription  of  the  genus. 


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Society  of  America  69: 1147-1153,  6  figs. 

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col.  pis.  In  von  der  Decken,  C.  C. ,  Reisen  in  Ost-Afrika  3  (2). 

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330  pp.,  52  pis.  Beijing. 


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57  f3):  98-112. 


212 


INDEX 

Index 


Invalid  names  are  in  italics;  references  to  descriptions  are  in  bold. 


abyssinica  194, 197,  201 

airensis  157, 159 

aliena  174 

alienus  175 

Amblypelta  155 

Ammocoris  191 

amphrysia  168, 170, 174 

angusta  179, 180 

an  gust  us  192 

annulipes  183, 184, 185, 187 

Anoplocerus  207 

Arenocoris  156, 191, 206, 207 

Atractus  191 

australis  166, 167, 194 

Austrocoris  155 

Bathysolen  206, 207 

bilineata  155 

Bothrostethus  207 

Boudicca  191 

breviceps  207 

brevitibialis  168, 170, 172, 174 

Brotheolus  154 

Brotheus  154 

brunneofusca  168, 170, 177, 178, 179 

calida  194, 196, 197, 198, 200 

campbelli  162, 163 

Ceraleptus  155, 206, 207 

Chariesterus  155 

Clavigralla  155, 190, 194, 206, 207, 

208 

Clavigralloides  207, 208 
clavipes  168, 170, 178 
concolor  182, 194 
conculata  163, 164 
Coriomeris  152, 206, 207, 209 
cornuta!81,182, 194 
Cristovallia  155 

delhiensis  157, 160, 161,162 
dispar  (Myla)  194, 198 
dispar  (Psilolomia)  169, 174 

egenus  191 
elevator  185 
elongata  206, 207 

fallen!  191, 192 


ferruginea  170, 172 

gestroi!91,192 
gracilis  194,  204 
graciloides  204 
Gralliclava  207,  208,  209 
granula  194, 198 
granulifera  187, 189 


Hoplolomia  152, 154, 156, 161, 163, 

205,207 
hoploxys 194, 197, 198, 200, 201, 

202,  204 

horrens  206,  208, 209 
hystrix  (Clavigralla)  187 
hystrix  (Mevanidea)  189, 190 

Indolomia  152, 154, 156, 163,  205, 

207 
intermedius  191, 192, 206, 208 

kilimana  189 

lamottei  170, 171, 172 
lata  (Myla)  193, 194, 200 
lata  (Psilolomia)  170, 179 
Iatissimusl91,192 
Loxocnemis  207 

Mevania  152, 187 

Mevanidea  156, 183, 185, 187, 207, 

208 

Mevaniella  187 
Mevaniomorpha  156, 183, 185, 189, 

207 

microphthalma  194, 202, 203, 204 
Microtelocerus  152,  206, 207 
Myla  152, 156, 165, 181, 193, 207, 

208 

Nemocoris  207 
Neohoplolomia  168 
Neomevaniomorpha  156, 183,  207 
nigeriensis  170, 171, 177, 208,  209 
nigrispina  193, 196 
niokensis  165, 193, 194, 201, 203 
numidianus  157, 159, 160 

Oncaspidia  207, 208 


onceroma  194, 203 

Paramyla  152, 154, 156, 165, 194, 

205,206,207 
parva  168, 170, 173, 174 
picta  185, 186, 187 
picta  186, 187 
Psammocoris  191 

Pseudomyla  156, 180, 183, 194,  207 
Pseudophloeus  191 
Psilocoris  168 
Psilolomia  156, 168, 183, 185, 207, 

208, 209 

pundaloyae  170, 177 
Pungral56, 179, 181,207 

quadrocephalus  157, 161 

Riptortus  154 

Risbecocoris  152, 154, 156, 162, 205, 

207,  208, 209 
rodhaini  183 

scabricula  161, 162, 163 
schnelli  194,  204, 205 
scutellaris  206, 207 
shadabi  207 
somalica  194,  200 
Spathocera  154 
spiniceps  187, 189, 190 
spinicollis  180, 181, 182, 183 
steeleae  170, 178 
Strobilotoma  207 
suspecta  165, 166, 194 

tomentosicollis  207 
tomentosus  156, 157, 159, 161 
Trallianus  155 
tuberculicollis  207 
typica  (Cristovallia)  155 
typica  (Neohoplolomia)  168, 172 

Ulmicola  207 
Urartucoris  206,  207 

Vilga  152, 154, 205,  207, 209 
vulgaris  170, 171, 175, 177, 178, 179 

waltli!91,192 
westwoodi  154 


Occasional  Papers  on  Systematic  Entomology 

New  Series 


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vast  literature  written  in  many  languages;  that  which  is  of  direct  economic  importance  and 
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A.  Watson  &  D.  T.  Goodger 

72pp.  inc.  4  colour  plates    27  February  1986 

No.  2.  An  annotated  checklist  of  the  Carabidae  (including  Cicindelinae,  Rhysodinae  and 

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N.  E.  Stork 

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Australian  ichneumonids  of  the  tribes  Labenini  and  Poecilocryptini 

By  I.  D.  Gauld  &  G.  A.  Holloway 

The  tribe  Pseudophloeini  (Hemiptera:  Coreidae)  in  the  Old  World  tropics  with  a  discussion  on 
the  distribution  of  the  Pseudophloeinae 

By  W.  R.  Dolling 

The  songs  of  the  western  European  grasshoppers  of  the  genus  Omocestusin  relation  to  their 
taxonomy  (Orthoptera:  Acrididae) 

By  D.  R.  Ragge 


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The  songs  of  the  western  European 
grasshoppers  of  the  genus  Omocestus  in 
relation  to  their  taxonomy  (Orthoptera: 
Acrididae) 


D.  R.  Ragge 


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Vol53  No  4  27  November  1986 


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Issued  27  November  1986 


The  songs  of  the  western  European  grasshoppers  of 
the  genus  Omocestus  in  relation  to  their  taxonomy 
(Orthoptera:  Acrididae) 

D.  R.  Ragge 

Department  of  Entomology,  British  Museum  (Natural  History),  Cromwell  Road,  London 
SW75BD 

Contents 

Synopsis 213 

Introduction 213 

Acknowledgements 214 

Abbreviations  of  depositories 214 

Methods 215 

Recording  and  analysing  the  songs 215 

Song  terminology 215 

Presentation 215 

The  song  as  a  taxonomic  character 219 

New  lectotype  designations 220 

Omocestus  Bolivar 221 

Keys  to  the  principal  western  European  species  of  Omocestus 222 

Key  based  primarily  on  morphological  characters 222 

Key  based  on  song  characters 224 

The  songs  of  the  western  European  Omocestus 226 

References .- 246 

Index 249 

Synopsis 

The  male  calling  songs  of  11  western  European  and  one  Canary  Island  species  of  the  Gomphocerine  genus 
Omocestus  are  described  and  fully  illustrated  with  oscillograms  at  three  or  four  different  speeds.  The 
courtship  song  is  also  described  for  the  six  species  in  which  such  a  song  is  known.  The  songs  of  six  species 
(panteli,  bolivari,  uhagonii,  minutissimus,  llorenteae  and  simony i}  are  described  for  the  first  time.  For  each 
species  a  full  list  is  provided  of  references  to  any  past  published  work  on  the  songs,  classified  according  to 
the  kind  of  information  given.  Notes  on  recognition,  using  both  morphology  and  song,  are  given  for  each 
species,  and  two  identification  keys  are  provided,  one  based  mainly  on  morphology  and  the  other  on  song. 
The  value  of  the  song  as  a  taxonomic  character  is  discussed. 

Introduction 

The  Gomphocerinae,  of  which  Omocestus  is  one  of  the  better  known  Old  World  genera,  include 
most  of  the  common  European  grasshoppers  of  open  grassland  and  moorland.  They  produce  the 
most  highly  developed  communicative  sounds  of  all  grasshoppers  and  are  widely  used  in 
bio-acoustic,  ecological  and  cytogenetic  research.  The  subfamily  includes  about  150  European 
species,  of  which  over  20  are  pests  in  parts  of  their  range,  seven  of  them  being  regarded  as  major 
ones  (Tsyplenkov,  1970). 

Omocestus  is  a  widespread  Palaearctic  genus,  including  some  40  species  living  in  habitats 
ranging  from  lush  lowland  meadow  to  steppe,  open  woodland  and  high  mountains.  Twenty 
species  have  been  recorded  from  western  Europe,  of  which  three,  rufipes,  petraeus  and 
particularly  haemorrhoidalis ,  become  pests  when  they  occur  in  sufficient  numbers,  causing 
damage  to  cereal  grasses,  hayfields,  alfalfa  and  other  cultivated  plants  in  western  and, 
especially,  eastern  Europe  (Bei-Bienko  &  Mishchenko,  1951;  Tsyplenkov,  1970). 

The  conspicuous  songs  of  Gomphocerine  grasshoppers  have  attracted  attention  for  centuries, 


Bull.  Br.  Mus.  not.  Hist.  (Ent.)  53  (4):  213-249  Issued  27  November  1986 


214  D.  R.  RAGGE 

but  it  is  only  since  the  pioneering  work  of  Faber  (1928)  that  they  have  been  considered  seriously 
as  an  aid  to  identification,  and  only  during  the  past  30  years  that  their  importance  in  taxonomic 
research,  and  particularly  in  resolving  complexes  of  sibling  species,  has  been  fully  recognized. 
This  importance  stems  from  the  current  belief  that  they  provide  the  main  mate  recognition 
system  in  these  insects  and  are  thus  able  to  maintain  reproductive  isolation  in  groups  of 
sympatric  species  that  would  otherwise  be  able  to  interbreed  freely.  Perdeck  (1957)  established 
experimentally  that  this  is  true  of  the  well-known  sibling  pair  Chorthippus  brunneus  (Thunberg) 
and  C.  biguttulus  (Linnaeus),  and,  although  comparable  studies  on  other  groups  of  Gom- 
phocerinae  are  still  greatly  needed,  it  seems  likely  that  such  studies  would  confirm  that  this 
function  of  the  song  is  widespread  in  the  subfamily. 

Grasshopper  songs  can  be  quite  easily  recorded  on  tape  and  their  rhythmic  patterns,  which 
provide  the  most  useful  taxonomic  information,  lend  themselves  well  to  oscillographic  analysis. 
It  is  thus  rather  surprising  that  there  has  so  far  been  no  comprehensive  account  of  these  songs, 
presented  group  by  group  and  illustrated  with  oscillograms.  The  main  aim  of  the  present  paper, 
and  of  similar  papers  in  preparation  on  other  groups  of  Orthoptera,  is  to  provide  such  an 
account,  arranged  systematically  by  group  and  fully  illustrated  with  oscillograms  at  several 
different  speeds.  It  is  hoped  that  this  form  of  presentation  will  make  the  information  most  useful 
in  identification  and  taxonomy,  as  well  as  providing  a  basic  reference  work  on  the  songs  of  these 
insects. 

No  information  has  previously  been  published  on  the  songs  of  six  of  the  12  species  included  in 
this  study  (panteli,  bolivari,  uhagonii,  minutissimus,  llorenteae  and  simonyi)  and  no  oscillograms 
have  previously  been  published  for  a  further  two  (petraeus  and  raymondi).  For  the  species  in 
which  published  information  on  the  song  is  already  available,  references  to  the  sources  are 
given,  classified  according  to  the  kind  of  information  published.  O.  simonyi,  known  only  from 
the  Canary  Islands,  is  not  strictly  within  the  scope  of  this  paper  but  is  included  for  convenience. 

The  song  is  still  unknown  in  the  following  nine  species  of  Omocestus  recorded  from  western 
Europe:  antigai,  burri,  corsicus,  femoralis,  kaestneri,  knipperi,  lopadusae,  navasi  and  uvarovi. 
These  species,  which  are  all  very  local  and  often  of  doubtful  status,  are  therefore  excluded  from 
this  study. 


Acknowledgements 

I  am  much  indebted  to  the  following,  who  have  kindly  lent  me  type-specimens  or  other  material  from  their 
respective  institutions:  Drs  R.  Danielsson,  M.  Donskoff,  P.  Grootaert,  K.  K.  Giinther,  A.  Kaltenbach,  V. 
Llorente,  F.  Pascual  and  J.  J.  Presa. 

My  thanks  are  also  due  to  Dr  N.  D.  Jago,  who  kindly  provided  me  with  tape  recordings  of  the  song  of  O. 
viridulus  made  by  him  in  the  French  Alps;  to  Mr  W.  G.  Tremewan,  who  was  kind  enough  to  collect  live 
males  of  O.  panteli  for  me  in  Teruel  Province,  Spain;  and  to  Mr  B.  Alexander,  who  kindly  brought  me  a  live 
male  of  O.  simonyi  from  Lanzarote,  Canary  Islands. 

I  am  particularly  grateful  to  my  colleague  Mr  W.  J.  Reynolds,  who  made  the  studio  recordings  used  in 
this  study,  helped  me  in  reviewing  the  past  literature  on  the  songs,  tested  the  identification  key  based  on 
morphological  characters  and  provided  valuable  comments  at  various  stages  in  the  preparation  of  the 
paper. 

Finally  I  wish  to  thank  my  wife,  who  has  given  me  invaluable  help  in  all  my  field-work. 


Abbreviations  of  depositories 

BMNH  British  Museum  (Natural  History),  London 

IRSNB  Institut  Royal  des  Sciences  Naturelles  de  Belgique,  Brussels 

MNCN  Museo  Nacional  de  Ciencias  Naturales,  Madrid 

MNHN  Museum  National  d'Histoire  Naturelle,  Paris 

MNHU  Museum  fur  Naturkunde  der  Humboldt-Universitat,  Berlin 

ZI  Zoologiska  Institution,  Lunds  Universitet,  Lund 


SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS  215 

Methods 

Recording  and  analysing  the  songs 

All  the  field  recordings  of  the  songs  used  in  this  study  were  made  while  the  insects  were  in  full 
sunshine  using  a  Uher  4000,  4200  or  4200IC  tape  recorder  and  AKG  D202  or  (recording  222/3 
only)  Shibaden  microphone;  the  tape  speed  was  always  19  cm/s.  The  subjects  were  approached 
stealthily  so  that  the  microphone  could  be  held  (or  rested  on  the  ground)  about  10  cm  from  them 
without  causing  any  disturbance. 

The  studio  recordings  were  made  in  the  BMNH  Acoustic  Laboratory  using  a  Kudelski  Nagra 
IV  tape  recorder  and  Sennheiser  MKH405  microphone;  the  tape  speed  was  19  or  38  cm/s.  In 
every  case  a  bench  lamp  was  used  to  provide  light  and  radiant  heat. 

Further  data  are  given  in  Tables  1  and  2  for  the  recordings  used  for  the  oscillograms 
reproduced  in  Figs  12-128.  However,  the  song  descriptions  given  for  each  species  are  in  most 
cases  based  on  many  more  recordings  of  numerous  songs,  all  of  which  were  analysed  oscillo- 
graphically.  The  oscillograms  reproduced  in  the  figures  were  chosen  as  being  typical  of  the 
species  concerned  and,  in  some  cases,  showing  the  extent  of  intraspecific  variation.  Although  I 
have  given  in  Tables  1  and  2  the  ambient  air  temperatures  at  the  time  of  recording,  it  is  really  the 
body  temperature  of  the  singing  insect  that  is  important,  and  this  is  more  dependent  on  radiant 
heat  than  on  the  air  temperature;  as  the  insects  were  always  receiving  radiant  heat  from  the  sun 
or  a  bench  lamp,  the  conditions  under  which  the  recordings  were  made  were  more  uniform  than 
the  different  air  temperatures  might  suggest. 

All  the  oscillograms  were  made  from  recorded  songs  with  a  Mingograf  34T  ink- jet  recorder. 

Song  terminology 

The  bio-acoustic  terms  used  are  defined  as  follows. 

Calling  song.  The  song  produced  by  an  isolated  male. 

Courtship  song.  The  special  song  produced  by  a  male  when  close  to  a  female. 

Syllable.  The  sound  produced  by  one  complete  up  and  down  movement  of  the  hind  legs 

(Fig- 1). 

Echeme.  A  first-order  assemblage  of  syllables  (Fig.  1). 
Echeme-sequence.  A  first-order  assemblage  of  echemes  (Fig.  1). 

Momentary  breaks  in  the  sound  (of  at  least  1-25  ms)  during  the  course  of  a  syllable  are 
referred  to  as  'gaps'. 

Presentation 

As  the  main  purpose  of  this  paper  is  to  give  information  on  songs,  I  have  not  included  either  a  full 
synonymy  or  a  formal  morphological  diagnosis  in  the  account  of  each  species.  I  have,  however, 
listed  references  to  all  past  descriptive  accounts  of  the  songs  (of  any  significance),  classified 
according  to  whether  they  include  oscillograms  (including  sound-level  tracings),  diagrams  (i.e. 
hand-drawn  representations  of  the  songs),  frequency  information,  musical  notation,  or  verbal 
description  without  any  of  these  additions;  any  commercially  available  disc  or  cassette  record- 
ings of  the  songs  are  also  listed.  These  references  are  not  intended  to  be  exhaustive  -  there  are 
many  brief  statements  about  the  songs,  especially  in  the  earlier  literature,  that  do  not  warrant 
inclusion;  my  aim  has  been  to  list  all  sources  that  the  reader  might  find  useful  to  refer  to  for 
additional  or  confirmatory  information  on  the  songs.  The  only  reference  I  have  included  to  a 
work  published  before  the  present  century  is  to  Yersin's  (1854)  short  but  admirable  account  of 
the  songs  of  38  European  species  of  Orthoptera  (including  two  of  Omocestus),  in  which  he 
attempted,  with  some  success,  to  represent  them  in  terms  of  musical  notation;  in  this  largely 
forgotten  work  he  demonstrated  clearly  for  the  first  time  the  striking  differences  between  the 
songs  of  the  three  closely  similar  species  Chorthippus  brunneus  (Thunberg),  C.  biguttulus 
(Linnaeus)  and  C.  mollis  (Charpentier). 
Notes  are  given  for  each  species  on  recognition  by  both  morphology  and  song,  and  these  are 


216 


D.  R.  RAGGE 


Table  1  Data  for  the  field  recordings  of  male  songs  of  Omocestus  used  for  oscillograms  reproduced  in  this 
study.  All  these  recordings  were  made  from  different  males.  Recordists'  names  are  abbreviated  as 
follows:  NDJ  =  N.  D.  Jago;  DRR  =  D.  R.  Ragge;  WJR  =  W.  J.  Reynolds. 


Species 


Locality 


Date  recorded  Shade  air          Recording 

and  collected       Recordist     temperature     No. 


viridulus 


rufipes 


haemorrhoidalis 


petraeus 


FRANCE:  Alpes- 
Maritimes,  near  St 
DalmasdeTende, 
Vallon  de  la  Miniere, 
1830m 

AUSTRIA:  Tyrol,  near 
Kufstein,  Kaisertal 
FRANCE:  Hautes- 
Pyrenees,  near 
Vielle-Aure,  Neste 
de  Couplan 

FRANCE:  Lozere,  near 
Mende,  Col  de 
Montmirat 
FRANCE:  Alpes- 
Maritimes,  near  St 
Martin  Vesubie,  Col  St 
Martin,  1400  m 

FRANCE:  Haute- Vienne, 
near  St 

Yrieix-la-Perche 
FRANCE:  Lozere, 
Causse  Mejean,  near 


26.viii.1977         NDJ 
13.viii.1973         DRR  24°C 


20.ix.1976  WJR 


14°C 


10.viii.1977         DRR  21°C 


26.viii.1977         DRR  22°C 


30.viii.1978         WJR  25°C 


222/3 
98/3 

183/2 

232/3 

241/11 
268/3 


Crosgarnon 

31.vii.1982 

DRR 

20°C 

482/5 

raymondi 

SPAIN:  Huesca,  8  km 
E.  of  Ainsa. 

3.ix.l978 

WJR 

31°C 

270/8 

SPAIN:  Madrid,  Puerto 

de  Galapagar,  800  m 
SPAIN:  Granada, 

17.  vi.  1984 

DRR 

30°C 

531/3 

Sierra  Nevada,  near 

Capileira,  1500  m 

22.  vi.  1984 

DRR 

24°C 

532/4 

panteli 

SPAIN:  Madrid,  near 
Navacerrada,  Valle 
de  Barranca 

27.vii.1983 

DRR 

22°C 

503/5 

SPAIN:  Granada, 

Sierra  Nevada,  Puerto 
de  la  Ragua,  2000  m 

l.viii.1983 

DRR 

25°C 

506/3 

broelemanni 

FRANCE:  Pyrenees- 
Orientales,  near 

Saillagouse,  Val 
d'Eyne 

15.ix.1978 

WJR 

23°C 

276/3 

bolivari 

SPAIN:  Granada, 

Sierra  Nevada,  Campos 
de  Otero,  2300  m 

31.vii.1983 

DRR 

21°C 

505/4 

SPAIN:  Granada, 

Sierra  Nevada,  Puerto 

de  la  Ragua,  2000  m 
Same  locality 

l.viii.1983 
l.viii.1983 

DRR 
DRR 

25°C 
25°C 

506/5 
507/2 

SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS 


217 


Table  1  -  cont. 


Species 


Locality 


Date  recorded  Shade  air          Recording 

and  collected       Recordist     temperature      No. 


uhagonii 


mmutissimus 


SPAIN:  Madrid, 
Sierra  de  Guadarrama, 
La  Bola  del  Mundo, 
2200m 
Same  locality 
Same  locality 

SPAIN:  Madrid, 
Sierra  de  Guadarrama, 
Puerto  de  los  Leones, 
1500m 


6.viii.l983 
6.viii.l983 
6.viii.l983 


DRR 
DRR 
DRR 


21°C 
21°C 

24°C 


5.viii.l983         DRR  23°C 


508/6 
509/1 
509/5 


508/3 


llorenteae                 SPAIN:  Granada, 

Sierra  Nevada,  near 

Dornajo,  1900m 
Same  locality 

30.vii.1983 
30.vii.1983 

DRR 
DRR 

27°C 
27°C 

504/6 
505/1 

Table  2  Data  for  the  studio  recordings  of  male  songs  of  Omocestus  used  for  oscillograms  reproduced  in 
this  study.  All  these  recordings  were  made  from  different  males  except  for  310  and  312,  which  were  made 
on  different  days  from  the  same  male  ofpanteli.  The  recordist  was  W.  J.  Reynolds  in  every  case. 


Species                     Locality 

Date 
collected 

Date 
recorded 

Air                    Recording 
temperature     No. 

viridulus                  ENGLAND:  Dorset, 

nearWareham, 

Morden  Bog 
ENGLAND:  Surrey, 
near  Mytchett  Lake 

18, 

22, 

vii.1976 
,  vii.1979 

21, 
26, 

vii.1976 
.vii.1979 

29°C 

32°C 

168 

307/2 

rufipes                     ENGLAND:  Kent, 
near  Goudhurst, 

Bedgebury  Forest 
ENGLAND:  Hampshire. 
New  Forest, 

18 

.vii.1976 

20 

.vii.1976 

26°C 

167/2 

Putties  Bridge 

4 

viii.1979 

6 

.viii.1979 

26°C 

319 

haemorrhoidalis 


FRANCE:  Pyrenees- 
Orientales,  near 
Saillagouse,  Val 
d'Eyne 


22.ix.1976 


29.ix.1976 


26°C 


177 


panteli 

SPAIN:  Valencia, 
near  Requena, 
700m 
Same  locality 
Same  locality 

27.vii.1979 
27.vii.1979 
27.vii.1979 

30.vii.1979 
31.  vii.1979 
l.viii.1979 

33°C 
30°C 
29°C 

310 
312 
315 

broelemanni 

FRANCE:  Pyrenees- 
Orientales,  near 

Saillagouse,  Val 
d'Eyne 

15.ix.1978 

19.ix.1978 

25°C 

266 

simonyi 

CANARY  ISLANDS: 
Lanzarote,  Teguise 

12.ix.1982 

15.ix.1982 

28°C 

562 

218 


D.  R.  RAGGE 


ECHEME 


•  HI  I  '  'I  1 1  it •  «  H  M 


|l»«**»|i|M 


SYLLABLE 


GAPS 


Omocesfus  raymonc/i 


ECHEME-SEQUENCE 


I         0-05  s 


•+-* 


•»HMH-H  >  I  » 


ECHEME 


0-05  s 


Omocesfus  pefraeus 


Fig.  1     Oscillograms  of  the  male  calling  songs  of  two  species  of  Omocestus,  showing  the  terminology  used 
in  this  paper. 


SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS  219 

followed  by  a  more  detailed  descriptive  account  of  the  calling  song  and,  when  known,  courtship 
song.  I  have  not  attempted  to  describe  any  further  kinds  of  sound  produced  by  the  males  (e.g. 
'rivalry  song',  'copulation  song')  as  these  are  not  stereotypically  patterned  and  seldom  of 
diagnostic  value.  Neither  have  I  considered  it  appropriate  to  describe  the  songs  produced  by 
females  when  in  a  sexually  receptive  state;  these  songs  are  seldom  heard  in  the  field  and  are  in 
any  case  broadly  similar  in  pattern  to  the  calling  songs  of  conspecific  males. 

The  songs  of  each  species  are  illustrated  by  oscillograms  at  three,  or  sometimes  four,  different 
speeds.  As  the  scale  lines  indicate,  the  three  speeds  given  for  all  species  are  (after  reduction  for 
printing)  10  mm/s,  80  mm/s  and  320  mm/s,  and  the  fourth  speed  included  for  the  songs  of  three 
species  (see  below)  is  1280  mm/s.  In  addition,  oscillograms  at  10  mm/s  of  typical  male  calling 
songs  of  all  12  species  are  shown  together  in  Figs  12-23  so  that  the  more  obvious  differences 
between  them  can  be  seen  at  a  glance. 

Grasshopper  songs  are  normally  produced  by  both  hind  legs  being  rubbed  simultaneously 
against  the  fore  wings,  and  oscillograms  of  such  'two-legged'  songs  show  the  sounds  produced  by 
the  movement  of  one  hind  leg  superimposed  on  those  produced  by  the  other.  The  two  hind  legs 
are  not  always  well  synchronized  and  in  some  species  follow  different  patterns  of  movement,  so 
that  there  is  sometimes  a  degree  of  'blurring'  of  the  oscillogram.  Moreover,  in  'two-legged' 
oscillograms  that  are  fast  enough  to  show  the  impacts  of  individual  stridulatory  pegs,  it  is 
impossible  to  determine  which  peg-impacts  are  produced  by  one  leg  since  these  are  mixed  with 
those  produced  by  the  other.  Although  these  points  are  not  particularly  relevant  to  the 
diagnostic  value  of  the  songs,  I  have  thought  it  worth  including  for  three  species  (viridulus, 
rufipes  and  panteli}  oscillograms  taken  from  the  calling  songs  of  males  with  only  one  hind  leg. 
The  difference  between  the  'one-legged'  and  'two-legged'  oscillograms  is  hardly  apparent  at  the 
slower  speeds,  but  becomes  more  obvious  at  320  mm/s  and  clearer  still  at  1280  mm/s.  The  highest 
speed  'one-legged'  oscillogram  of  panteli  (Fig.  89)  shows  the  individual  peg-impacts  with 
particular  clarity. 

The  song  as  a  taxonomic  character 

Recognition  of  the  diagnostic  value  of  the  songs  of  Orthoptera  may  be  said  to  have  begun  in  the 
mid-nineteenth  century,  when  Fischer  (1849,  1850,  1853)  published  a  number  of  song  descrip- 
tions in  primarily  faunal  or  systematic  works,  and  Yersin  (1852,  1853  and  especially  1854) 
produced  a  series  of  papers  specifically  on  the  songs  and  their  value  in  identification.  Yersin's 
work  of  1854,  in  which  he  described  the  songs  of  38  European  species  of  Orthoptera,  was 
particularly  noteworthy  in  that  it  included  the  first  attempt  to  illustrate  the  songs  graphically,  in 
the  form  of  musical  notation. 

Little  further  progress  was  made  in  exploiting  the  songs  in  taxonomy  until  Faber's  work  of 
1928,  in  which  he  gave,  for  the  first  time,  an  identification  key  to  the  German  Orthoptera  based 
on  their  songs.  Many  German  studies  on  the  songs  of  the  European  Orthoptera  followed, 
notably  by  Faber  (especially  1929, 1932  and  1936)  and  Jacobs  (especially  1950),  culminating  in 
two  major  works  published  almost  simultaneously  (Faber,  1953;  Jacobs,  1953).  Most  of  this 
work  was  concerned  with  the  detailed  description  of  the  songs  and  associated  behaviour,  and 
there  was  still  no  serious  attempt  to  use  the  songs  in  taxonomic  research. 

Jacobs  (1950,  1953)  used  simple  diagrams  against  a  time  scale  to  illustrate  the  rhythmic 
patterns  of  the  songs.  Ragge  (1965)  introduced  a  more  sophisticated  kind  of  diagram,  amounting 
to  a  simplified  oscillogram,  and  diagrams  of  this  kind  have  since  been  used  by  Hoist  (1970), 
Luquet  (1978),  Wallin  (1979)  and,  in  a  slightly  different  form,  by  Duijm  &  Kruseman  (1983)  and 
Bellmann  (1985).  Identification  keys  based  on  the  songs  were  provided,  for  restricted  parts  of 
Europe,  by  Luquet  (the  Mont  Ventoux  area  of  the  French  Alps,  Acrididae  only),  Wallin 
(Sweden),  Duijm  &  Kruseman  (Benelux)  and  Bellmann  (West  Germany).  All  these  studies  are 
concerned  with  regional  faunas  and  contain  no  taxonomic  research,  although  Luquet  gives  much 
emphasis  to  the  importance  of  the  songs  in  taxonomy. 

The  first  serious  application  of  the  songs  of  Orthoptera  to  their  taxonomy  was  in  work  on  the 
North  American  crickets  (Gryllidae).  Following  the  pioneering  work  of  Fulton  (1931,  1952), 


220  D.  R.  RAGGE 

Alexander  (1957)  and  Alexander  &  Thomas  (1959)  reviewed  the  taxonomy  of  North  American 
species  of  Acheta  and  Nemobius,  and  Walker  (1962,  1963)  carried  out  similar  studies  on 
Oecanthinae;  in  all  these  studies  the  songs  played  a  crucial  part  in  resolving  the  taxonomy  of 
morphologically  similar  species.  Similar  work  on  other  groups  followed,  some  of  it  reviewed  by 
Alexander  (1962)  and  Walker  (1964).  In  1972  Alexander,  Pace  &  Otte  produced  an  account  of 
the  'singing  insects'  of  Michigan  in  which  they  gave  keys  based  mainly  on  the  songs  for  all  the 
Ensifera. 

More  recent  taxonomic  studies  on  Orthoptera  in  which  the  songs  have  played  a  significant  part 
include  those  of  Bailey  (1975,  1980)  on  African  and  Australian  Copiphorine  Tettigoniidae, 
Walker  &  Greenfield  (1983)  on  Caribbean  Neoconocephalus  (Tettigoniidae),  Otte  &  Alexander 

(1983)  on  Australian  Gryllidae,  Otte  &  Cade  (1983  and  later)  on  African  Gryllidae,  Heller 

(1984)  on  Poecilimon  (Tettigoniidae)  and  Ragge  &  Reynolds  (1984)  on  western  European 
Euchorthippus  (Acrididae). 

As  can  be  seen  at  a  glance  from  Figs  12-23,  the  male  calling  songs  of  the  western  European 
species  of  Omocestus  provide  most  useful  taxonomic  characters.  Recorded  songs,  with 
appropriate  oscillographic  analyses,  can  be  used  to  identify  the  species  even  if  no  specimens  are 
available  for  morphological  study  (see  the  key  on  p.  224). 

The  calling  songs  of  the  12  species  embraced  by  the  present  study  fall  into  two  clearly  different 
kinds:  the  rapid  sequence  of  echemes  produced  by  petraeus,  minutissimus  and  llorenteae,  and 
the  single  or  widely  separated  echemes  produced  by  the  remaining  species.  As  would  be 
expected,  the  rapidly  repeated  echemes  of  petraeus,  minutissimus  and  llorenteae  are  composed 
of  very  rapidly  repeated  syllables,  almost  always  more  than  80/s,  whereas  the  syllable  repetition 
rate  of  the  remaining  species  is  almost  always  less  than  40/s.  Within  this  trio  of  species  petraeus 
stands  a  little  apart  from  the  other  two  in  having  even  shorter  and  more  rapidly  repeated 
echemes,  but  the  songs  of  minutissimus  and  llorenteae  are  broadly  similar  to  each  other  in 
pattern,  differing  mainly  in  the  duration  of  the  echeme-sequence  and  the  number  of  echemes  of 
which  it  is  composed. 

Among  the  remaining  species,  viridulus  and  rufipes  have  much  longer  echemes  than  all  the 
others;  they  are  very  similar  to  each  other  in  both  calling  and  courtship  songs,  differing  mainly  in 
the  duration  of  the  calling  song  echeme.  The  songs  of  all  the  other  species  consist  of  relatively 
short  echemes  distinguished  from  one  another  by  duration,  syllable  repetition  rate,  and  number 
and  distribution  of  gaps.  The  song  differences  shown  by  the  four  brachypterous  species  bolivari, 
uhagonii,  minutissimus  and  llorenteae  are  particularly  useful  for  field  identification;  two  or  three 
of  these  species  sometimes  occur  together  in  the  same  locality  and  are  sufficiently  similar  in 
appearance  to  be  quite  easily  confused  with  one  another. 

Extreme  caution  has  to  be  used  in  drawing  phylogenetic  conclusions  from  resemblances  or 
differences  in  grasshopper  songs,  since  it  is  well  known  from  studies  on  the  Chorthippus 
biguttulus  group  that  apparently  closely  related  species,  capable  of  producing  vigorous  and 
fertile  hybrids,  can  have  strikingly  different  songs.  From  the  known  songs  of  Omocestus  I  am 
inclined  to  make  such  inferences  in  only  two  cases  of  song  resemblance.  The  first  is  the 
well-known  pair  viridulus  and  rufipes,  in  which  the  strong  resemblance  in  both  calling  and 
courtship  songs  suggests  that  they  are  sister  species.  The  second  is  the  pair  of  brachypterous, 
montane  species  minutissimus  and  llorenteae,  in  which  the  close  resemblance  in  calling  song  (as 
well  as  morphology)  also  suggests  a  comparatively  recent  common  ancestry. 


New  lectotype  designations 

In  the  course  of  examining  type-material  I  have  taken  the  opportunity  of  designating  lectotypes 
for  the  following  species  included  in  this  study. 

Gryllus rufipes  Zetterstedt,  1821:  90. 

Following  Ander  (1943: 10)  I  am  regarding  only  two  adult  males  from  Zetterstedt's  original  material  (in  the 

ZI,  Lund)  as  being  eligible  as  syntypes,  and  I  have  selected  and  labelled  one  of  these,  bearing  Zetterstedt's 


SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS  221 

ochre-yellow  label  (indicating  Ostergotland),  as  lectotype.  I  have  labelled  the  other  eligible  adult  male, 
which  bears  no  colour-coded  label,  as  a  paralectotype. 

Acridium petraeum  Brisout,  1855:  cxiv. 

There  is  in  the  IRSNB,  Brussels  a  long  series  of  both  sexes  of  this  species,  all  labelled  'Lardy',  the 
type-locality.  These  specimens  are  also  labelled  as  being  from  the  collection  of  de  Selys  Longchamps,  who 
had  earlier  acquired  Brisout's  collection,  and  it  seems  almost  certain  that  they  include  at  least  part  of 
Brisout's  type-series.  I  have  selected  and  labelled  a  male  bearing  the  handwritten  label  'Lardy  pres  de 
Paris'  as  lectotype.  As  there  are  many  other  specimens  of  this  and  various  other  species  of  Omocestus  (and 
other  genera)  in  the  de  Selys  Longchamps  Collection  labelled  'Lardy',  I  have  not  attempted  to  determine 
which  of  these  are  also  likely  to  belong  to  Brisout's  type-series  and  so  have  labelled  no  specimens  as 
paralectotypes. 

Stenobothrus  Brolemanni  Azam,  1906: 128. 

Dr  M.  Donskoff  of  the  MNHN,  Paris  has  kindly  sent  me  the  male  from  Azam's  type-series  that  seems  most 

eligible  as  a  lectotype  and  I  have  so-labelled  it. 

Omocestus  BolivariChopard,  1939: 172. 

The  type-series  of  this  species  (in  the  MNHN,  Paris)  consists  of  one  male  and  one  female,  and  I  have 
selected  and  labelled  the  male  as  lectotype.  The  altitude  given  in  the  original  description,  '2000  m. 
environ',  clashes  with  the  '3000  m'  given  on  the  locality  label,  which  I  think  is  more  likely  to  be  accurate.  I 
have  labelled  the  female  as  a  paralectotype. 

Gomphocerus  (Stenobothrus)  Uhagonii  Bolivar,  1876:  324. 

Dr  J.  J.  Presa  of  the  Universidad  de  Murcia  very  kindly  delivered  to  me  by  hand  a  male  syntype  of  this 
species  (from  the  MNCN,  Madrid)  labelled  as  being  from  Navarredonda,  one  of  the  type-localities,  and  I 
have  labelled  it  as  lectotype. 

OMOCESTUS  Bolivar 

Omocestus  Bolivar,  1878: 427  [as  subgenus  of  Gomphocerus  Thunberg];  Burr,  1904:  320  [raised  to  genus]. 

Type-species:  Gryllus  Locusta  viridulus  Linnaeus,  by  subsequent  designation  (Kirby,  1910: 172). 
Dirshius  Harz,  1975:  710  [as  subgenus  of  Omocestus  Bolivar].  Type-species:  Gryllus  haemorrhoidalis 

Charpentier,  by  original  designation.  Syn.  n. 

DIAGNOSIS,  cf  $•  Head  of  typical  Gomphocerine  shape,  foveolae  well  developed.  Antennae  not  clubbed. 
Pronotal  lateral  carinae  varying  from  sharply  incurved  to  almost  straight.  Brachypterous  to  macropterous. 
Fore  wings  without  bulge  on  anterior  margin  of  precostal  area;  medial  area  not  conspicuously  widened 
(except  in  O.  uvarovi  Zanon ,  whose  generic  assignment  is  uncertain) .  Hind  wings  varying  from  transparent 
to  strongly  smoky.  Ovipositor  varying  in  length,  normally  without  lateral  teeth  but  sometimes  showing 
tendency  towards  their  development  (e.g.  O.  broelemanni).  Tympanal  aperture  slit-like. 

DISCUSSION.  Omocestus  can  be  defined  only  on  negative  characters:  it  lacks  the  precostal  bulge  of 
Chorthippus  Fieber,  the  toothed  ovipositor  of  Stenobothrus  Fischer,  and  the  clubbed  antennae  of 
Gomphocerus  Thunberg,  Gomphocerippus  Roberts  and  Myrmeleotettix  Bolivar.  Like  several  other  genera 
in  the  large  and  rather  intractable  subfamily  Gomphocerinae,  it  is  no  more  than  a  group  of  convenience  and 
is  very  unlikely  to  be  holophyletic.  Harz  (1975)  attempted  to  split  it  into  two  subgenera  on  the  basis  of  the 
degree  of  curvature  of  the  pronotal  lateral  carinae,  but  the  species  show  every  gradation  between  sharply 
incurved  and  almost  straight  carinae,  and  it  is  thus  impossible  to  draw  a  clear  dividing  line  between  two 
subgroups. 

Jago  (1971)  treated  Omocestus  as  a  subgenus  of  Stenobothrus,  thus  reverting  to  the  status  accorded  to  it 
by  Bolivar  (1897)  and  Yakobson  &  Bianki  (1902).  Jago's  study  was  based  entirely  on  males,  however,  and 
so  did  not  take  into  account  the  toothed  ovipositor  as  the  main  diagnostic  character  of  Stenobothrus  s.  str.  I 
prefer  to  follow  Harz  (1975)  and  other  recent  workers  on  the  European  fauna  in  regarding  this  character  as 
sufficient  for  a  generic  distinction.  Jago  also  regarded  Kirby 's  designation  of  viridulus  as  the  type-species  of 
Omocestus  as  invalid  on  the  grounds  that,  at  the  time  of  the  original  description  of  the  genus,  this  species 
was  listed  by  Bolivar  (1878:  460)  under  Chorthippus  rather  than  Omocestus.  However,  on  p.  427  of 
Bolivar's  work,  where  Omocestus  is  first  established  and  diagnosed,  viridulus  is  clearly  included  in  it;  the 
heading  'Chorthippus'  in  the  list  on  p.  460  has  clearly  been  inadvertently  misplaced,  appearing  above 
rufipes  instead  of  in  its  correct  position  above  pullus.  I  am  therefore  accepting  Kirby 's  type-species 
designation  as  valid. 

The  male  callings  songs  show  a  wide  range  of  patterns  in  Omocestus,  varying  from  the  prolonged  single 


222  D.  R.  RAGGE 

echemes  of  viridulus  and  rufipes  to  the  rapid  sequence  of  short  echemes  oipetraeus.  This  is  not  in  itself  an 
indication  of  polyphyly,  however,  as  some  complexes  of  sibling  grasshopper  species  that  are  capable  of 
interbreeding  (e.g.  the  Chorthippus  biguttulus  group)  show  equally  striking  differences  in  male  calling 
song. 

DISTRIBUTION.  All  Europe,  the  larger  Mediterranean  islands,  North  Africa  and  temperate  Asia  as  far  as 
China. 

INCLUDED  SPECIES.  O.  alluaudi  Uvarov,  O.  antigai  (Bolivar),  O.  aymonissabaudiae  Salfi,  O.  bolivari 
Chopard,  O.  broelemanni  (Azam),  O.  burri  Uvarov,  O.  caucasicus  Tarbinskii,  O.  corsicus  Chopard,  O. 
cuonaensis  Yin,  O.  demokidovi  Ramme,  O.  enitor  Uvarov,  O.  femoralis  Bolivar,  O.  haemorrhoidalis 
(Charpentier),  O.  heymonsi  (Ramme),  O.  hingstoni  Uvarov,  O..  kaestneri  (Harz),  O.  knipperi  Harz,  O. 
lecerfi  Chopard,  O.  lepineyi  Chopard,  O.  llorenteae  Pascual,  O.  lopadusae  La  Greca,  O.  lucasii  (Brisout), 
O.  minutissimus  (Bolivar),  O.  minutus  (Brulle),  O.  megaoculus  Yin,  O.  motuoensis  Yin,  O.  nanus Uvarov, 
O.  navasi Bolivar,  O.  nyalamus  Xia,  O.  panteli  (Bolivar),  O.  petraeus  (Brisout),  O.  raymondi  (Yersin),  O. 
rufipes  (Zetterstedt),  O.  simony i  (Krauss),  O.  tibetanus  Uvarov,  O.  tzendsureni  Giinther,  O.  uhagonii 
(Bolivar),  O.  uvarovi  Zanon,  O.  viridulus  (Linnaeus),  O.  znojkoi  Mishchenko. 

Keys  to  the  principal  western  European  species  of  Omocestus 

The  only  western  European  species  not  included  in  these  keys  are  very  local  forms,  usually  of 
doubtful  status,  in  which  the  song  is  so  far  unknown. 

Two  keys  are  provided  here,  one  based  on  morphological  characters  but  with  notes  on  song 
differences  added  where  appropriate,  and  one  based  entirely  on  the  male  calling  songs.  Some  of 
the  species  are  very  difficult  to  separate  morphologically  and,  in  the  absence  of  song  informa- 
tion, some  specimens  may  not  be  correctly  identified  using  the  morphological  key.  Reference 
should  in  any  case  always  be  made  to  the  notes  given  on  recognition  in  the  accounts  of  each 
species. 

Key  based  primarily  on  morphological  characters 

This  key  is  for  sexually  mature  adults,  either  freshly  collected  or  showing  good  colour  preservation. 
O.  simony  i,  which  is  known  only  from  the  Canary  Islands,  is  not  included. 

1  Rather  brachypterous ,  fore  wings  less  than  3  times  length  of  pronotum 

-  Fully  winged,  fore  wings  more  than  3  times  length  of  pronotum 10 

2  Larger:  length  of  hind  femur  more  than  9-2  mm  in  cf,  more  than  11-0  mm  in  $.  (Eastern 

Pyrenees  and  Catalonia) broelemanni  (p.  236) 

Smaller:  length  of  hind  femur  less  than  9-2  mm  in  cf,  less  than  11  -Omm  in  $.  (Not  known  from 
Pyrenees) 3 

3  Fore  wings  not  or  hardly  projecting  beyond  hind  wings  (when  flexed);  medial  area  with  large 

dark  spots 4 

Fore  wings  pro j  ecting  well  beyond  hind  wings  (when  flexed) ;  medial  area  without  spots 5 

4  Fore  wings  more  than  2-5  times  length  of  pronotum  in  cf ,  more  than  2-1  times  in  9 

llorenteae  (p.  243) 
Fore  wings  less  than  2-5  times  length  of  pronotum  in  cf ,  less  than  2-1  times  in  $ 

minutissimus  (p.  243) 

5  Male 6 

Female 

6  Cerci  laterally  compressed  towards  tip uhagonii  (p.  237) 

Cerci  simply  conical,  not  laterally  compressed  towards  tip 

7  Hind  wings  reaching  less  than  halfway  along  fore  wings  (when  flexed).  Length  of  pronotum 

usually  more  than  2-5  mm.  (Song  a  single  echeme,  as  in  Fig.  19) bolivari  (p.  237) 

-  Hind  wings  reaching  more  than  halfway  along  fore  wings  (when  flexed).  Length  of  pronotum 

usually  less  than  2-5  mm.  (Song  a  sequence  of  echemes,  as  in  Fig.  21) minutissimus  (p.  243) 

8  Fore  wings  less  than  1-3  times  length  of  pronotum bolivari  (p.  237) 

Fore  wings  more  than  1  -4  times  length  of  pronotum 9 


SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS 


223 


ruf 


uho 


5  mm 


roy 


10 


pan 
11 


Figs  2-11  Ovipositors  and  pronota  of  species  of  Omocestus.  2-5.  Lateral  view  of  the  ovipositor  of  (2)  O. 
viridulus,  (3)  O.  rufipes,  (4)  O.  uhagonii,  (5)  O.  minutissimus.  6-11.  Dorsal  view  of  the  pronotum  of  (6) 
O.  viridulus,  (7)  O.  rufipes,  (8)  O.  haemorrhoidalis ,  (9)  O.  petraeus,  (10)  O.  raymondi,  (11)  O.  panteli. 


9    Ovipositor  shaped  as  in  Fig.  5,  lower  valves  with  strongly  sigmoid  ventral  profile 

minutissimus  (p.  243) 

-  Ovipositor  shaped  as  in  Fig.  4,  lower  valves  with  weakly  sigmoid  ventral  profile uhagonii  (p.  237) 

10  Pronotal  lateral  carinae  straight  or  almost  so  in  prozona  (Fig.  11).  (Iberian  Peninsula) 

panteli  (p.  236) 

-  Pronotal  lateral  carinae  distinctly  incurved  or  angled  in  prozona  (not  as  in  Fig.  11) 11 

11  Fore  wings  falling  well  short  of  hind  knees,  their  length  less  than  7-4  mm  in  cf ,  less  than  8-0  mm 

in  $ .  (Mountains in  S.  Spain) Ilorenteae (p.  243) 

-  Fore  wings  usually  reaching  at  least  to  hind  knees,  their  length  more  than  7-4  mm  in  d" ,  more 

than8-0mmin$ 12 

12  Hind  wings  completely  transparent  (except  sometimes  for  part  of  the  subcostal  area) .  Pronotal 

lateral  carinae  as  in  Fig.  9.  (Song  a  rapid  sequence  of  echemes,  as  in  Fig.  15) petraeus  (p.  233) 


224  D.  R.  RAGGE 

Hind  wings  at  least  slightly  smoky  towards  tip.  Pronotal  lateral  carinae  as  in  Figs  6,  7,  8  or  10. 
(Song  a  single  echeme ,  repeated  after  an  interval  of  at  least  several  seconds) 13 

13  Male 14 

Female 17 

14  Abdomen  with  no  orange  or  red  colouring.  (Song  echeme  lasting  more  than  12  s). .     viridulus(p.  226) 
Abdomen  with  at  least  some  orange  or  red  colouring.  (Song  echeme  lasting  less  than  12s) 15 

15  Abdomen  with  at  least  the  distal  5  sternites  coloured  red  or  orange-red.  Pronotal  lateral 

carinae  gently  incurved  in  prozona,  as  in  Fig.  7.  (Song  echeme  lasting  more  than  4  s) 

rufipes(p.  227) 

Abdomen  without  reddish  colouring  on  sternites  or  with  such  colouring  restricted  to  the  distal  3 
or  4  sternites.  Pronotal  lateral  carinae  more  strongly  incurved  in  prozona,  as  in  Figs  8  or  10. 
(Song  echeme  lasting  less  than  4s) 16 

16  Fore  wings  reaching  well  beyond  hind  knees,  more  than  4  times  length  of  pronotum.  (Song 

echeme  lasting  less  than  2s) raymondi(p.  233) 

Fore  wings  not  or  hardly  reaching  hind  knees,  less  than  4  times  length  of  pronotum.  (Song 
echeme  lasting  more  than  2  s) haemorrhoidalis(p.  227) 

17  Ovipositor  long,  as  in  Fig.  2.  Sides  of  body  often  green  (but  sometimes  brown  or  reddish) 

viridulus(p.  226) 

-  Ovipositor  shorter,  similar  in  length  to  Fig.  3.  Sides  of  body  never  green 18 

18  Abdomen  with  at  least  the  more  distal  sternites  coloured  red  or  orange-red.  Pronotal  lateral 

carinae  gently  incurved  in  prozona,  as  in  Fig.  7 rufipes(p.  227) 

-  Abdominal  sternites  with  no  reddish  colouring.  Pronotal  lateral  carinae  more  strongly  in- 

curved in  prozona,  as  in  Figs  8  or  10 19 

19  Fore  wings  reaching  beyond  hind  knees,  more  than  4  times  length  of  pronotum  . . .      raymondi  (p.  233) 

-  Fore  wings  not  or  hardly  reaching  beyond  hind  knees,  less  than  4  times  length  of  pronotum 

haemorrhoidatis(p.  227) 


Key  based  on  song  characters  (see  especially  Figs  12-23) 

This  key  is  based  on  the  calling  songs  produced  by  isolated  males  in  warm,  sunny  conditions.  As  far  as 
couplet  5  the  song  differences  can  be  easily  detected  by  the  human  ear  assisted  by  the  second  hand  (or 
digital  count)  of  a  watch.  From  couplet  6  onwards  some  of  the  differences  require  oscillographic  analysis, 
or  at  least  the  ability  to  play  back  the  recorded  song  at  a  slower  tape  speed. 
See  p.  00  for  definitions  of  the  terms  'echeme',  'syllable'  and  'gap'. 

1  Song  a  series  of  echemes  in  rapid  succession  (at  least  1/s) 2 

-  Song  a  single  echeme  or  a  series  of  echemes  separated  by  intervals  of  at  least  3s 4 

2  Echemes  repeated  at  the  rate  of  at  least  3/s  (Figs  55-63) petraeus(p.  233) 

Echemes  repeated  at  the  rate  of  about  2/s 3 

3  Song  consisting  of  about  1 1-15  echemes  and  lasting  about  6-8  s  (Figs  123-128) ....     llorenteae  (p.  243) 

-  Song  consisting  of  about  6-10  echemes  and  lasting  about  3-5  s  (Figs  117-1 19)       minutissimus(p.  243) 

4  Song  consisting  of  a  single  echeme  lasting  more  than  4s 5 

Song  consisting  of  one  or  more  echemes,  each  lasting  less  than  4s 6 

5  Echeme  lasting  more  than  12  s  (Figs  24-33) viridulus  (p.  226) 

Echeme  lasting  less  than  12s  (Figs  34-45) rufipes(p.  227) 

6  Syllable  repetition  rate  less  than  10/s 

Syllable  repetition  rate  more  than  10/s 8 

7  Each  syllable  with  more  than  7  gaps  (Figs  108-1 16).  (Iberian  Peninsula) uhagonii(p.231) 

-  Each  syllable  with  fewer  than  6  gaps  (Figs  120-122).  (Canary  Islands) simonyi  (p.  246) 

8  Syllable  repetition  rate  less  than  20/s 9 

-  Syllable  repetition  rate  more  than  20/s 11 

9  Echeme  lasting  more  than  2  s  (Figs  90-98) broelemanni(p.  236) 

-  Echeme  lasting  less  than  2s  (Figs  64-72  or  99-107) 10 


SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS 


"I 
O 


00 

c 


(N 


CN 


•O 


s 


K 


00 


cs 


o 

O) 

o 


O 
CN 


226  D.  R.  RAGGE 

10  Gaps  becoming  obscured  towards  the  end  of  the  echeme  (Figs  64-72).  (Widespread  in  Iberian 

Peninsula,  S.  France,  NW.  Italy  and  North  Africa) raymondi(p.  233) 

Gaps  persisting  to  the  end  of  the  echeme  (Figs  99-107).  (Known  only  from  the  higher  parts  of 
the  Sierra  Nevada  in  S.  Spain) bolivari(p.  237) 

1 1  Echeme  lasting  more  than  2  s  (Figs  46-54) haemorrhoidalis(p.  227) 

-    Echeme  lasting  less  than  2  s  (Figs  73-89) panteli(p.  236) 


The  songs  of  the  western  European  Omocestus 

Omocestus  viridulus  (Linnaeus) 

(Figs  2,  6,  24-33) 

Gryllus  Locusta  viridulus  Linnaeus,  1758:  433.  Lectotype  cf,  SWEDEN:  Gotland,  Burgsvik  (Linnean 
Society  of  London),  designated  by  Marshall  (1983:  394)  [examined]. 

REFERENCES  TO  SONG.  Oscillogram:  Eisner,  1974,  1975;  Grein,  1984;  Haskell,  1957,  1958;  Kutsch,  1976; 
Kutsch  &  Schiolten,  1979;  Loher  &  Broughton,  1955;  Zhantiev,  1981.  Diagram:  Bellmann,  1985a;  Duijm 
&  Kruseman,  1983;  Haskell,  1957;  Hoist,  1970;  Jacobs,  1950, 1953;  Ragge,  1965;  Wallin,  1979.  Frequency 
information:  Haskell,  1957,  1958.  Musical  notation:  Yersin,  1854.  Verbal  description  only:  Beier,  1956; 
Faber,  1928,  1953;  Haskell,  1955;  Harz,  1957;  Skovmand  &  Pedersen,  1978,  1983;  Weber,  1984;  Weih, 
1951.  Disc  recording:  Grein,  1984;  Ragge,  Burton  &  Wade,  1965.  Cassette  recording:  Bellmann,  19856; 
Wallin,  1979. 

RECOGNITION.  Males  of  this  common  species  may  be  easily  distinguished  from  the  rather  similar  species 
rufipes  and  haemorrhoidalis  by  the  lack  of  any  red  colouring  on  the  abdomen,  and  females  by  the 
conspicuously  longer  ovipositor  (Fig.  2).  Both  sexes  also  lack  the  dark  markings  in  the  medial  area  of  the 
fore  wings  that  are  normally  present  in  the  other  two  species,  and  the  pronotal  lateral  carinae  are  usually 
much  less  incurved  in  the  prozona  (Fig.  6). 

In  the  field  the  conspicuously  loud  calling  song  of  the  male  provides  an  easy  means  of  identifying  this 
species.  Although  similar  in  the  quality  of  sound  to  the  songs  of  rufipes  and  haemorrhoidalis,  the  song  of 
viridulus  is  more  prolonged,  usually  lasting  for  more  than  12  s. 

SONG  (Figs  24-33).  The  calling  song  is  an  echeme  usually  lasting  about  12-25  s  and  consisting  of 
syllables  repeated  at  the  rate  of  about  15-20/s.  The  echeme  beings  quietly  (the  first  few  leg 
movements  producing  no  audible  sound)  and  gradually  increases  in  loudness  until  the  maximum 
intensity  is  reached  after  a  few  seconds;  the  echeme  then  continues  at  a  constant  intensity  until 
reaching  an  abrupt  end.  The  syllable  repetition  rate  is  highest  at  the  beginning  of  the  echeme 
(usually  18-20/s),  gradually  lessening  towards  the  end  (when  it  is  usually  15-18/s).  The  song  is  a 
conspicuous  summer  sound  in  much  of  the  European  countryside,  louder  than  the  songs  of  most 
other  common  grasshoppers. 

In  the  presence  of  a  female  the  male  produces  longer  echemes,  usually  lasting  more  than  30  s 
and  occasionally  more  than  a  minute;  one  of  the  hind  legs  is  moved  through  a  noticeably  wider 
angle  than  the  other  and  produces  most  of  the  sound.  After  a  series  of  these  echemes  with  short 
pauses  (about  10-15  s)  between  them,  there  is  a  quite  different  and  much  quieter  echeme  lasting 
about  3-5  s  and  composed  of  syllables  repeated  at  the  rate  of  about  12-16/s  (Fig.  26).  This  is 
normally  followed  by  a  series  of  loud  syllables  and  an  attempt  to  copulate  with  the  female.  If  this 
attempt  is  unsuccessful  the  male  will  usually  produce  a  series  of  sharp  'ticks'  (Fig.  27)  by  kicking 
backwards  with  the  hind  tibiae  (as  in  the  calling  song  of  Stethophyma  grossum  (Linnaeus)) 
before  beginning  another  sequence  of  courtship  echemes.  Usually  the  two  hind  tibiae  are  kicked 
simultaneously,  but  sometimes  one  at  a  time,  changing  haphazardly  (rarely  regularly)  from  one 
side  to  the  other.  The  number  of  ticks  is  very  variable  but  is  usually  between  5  and  15,  and  the 
repetition  rate  is  rather  irregular,  generally  about  1-2/s. 

DISTRIBUTION.  This  common  species  occurs  in  fairly  moist  habitats  throughout  Europe,  except  for  the 
extreme  north  and  the  southern  parts  of  the  Iberian,  Italian  and  Balkan  peninsulas;  its  range  extends 
eastwards  to  Siberia  and  Mongolia. 


SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS  227 

Omocestus  rufipes  (Zetterstedt) 
(Figs  3,  7,  34-45) 

Gryllus  rufipes  Zetterstedt,  1821:  90.  LECTOTYPE  cf ,  SWEDEN:  Ostergotland,  Larketorp  (ZI,  Lund), 
here  designated  (see  p.  220)  [examined].  [Validity  established  by  von  Borck  (1848: 124)  under  Article  24 
of  the  International  Code  of  Zoological  Nomenclature.} 

Gryllus  ventralis  Zetterstedt,  1821:  89.  Holotype  $,  SWEDEN:  Skane,  near  Tranas,  Asperod  ['Esperod'] 
(ZI,  Lund)  [examined]. 

REFERENCES  TO  SONG.  Oscillogram:  Broughton,  1955;  Grein,  1984;  Loher  &  Broughton,  1955;  Schmidt  & 
Baumgarten,  1977.  Diagram:  Bellmann,  1985a;  Duijm  &  Kruseman,  1983;  Hoist,  1970;  Jacobs,  1950, 
1953;  Luquet,  1978;  Ragge,  1965;  Wallin,  1979.  Frequency  information:  Busnel,  1955;  Loher  &  Brought- 
on, 1955.  Musical  notation:  Yersin,  1854.  Verbal  description  only:  Beier,  1956;  Chopard,  1922;  Faber, 
1953;  Harz,  1957;  Weih,  1951.  Disc  recording:  Andrieu  &  Dumortier,  1963;  Grein,  1984;  Ragge,  Burton  & 
Wade,  1965.  Cassette  recording:  Bellmann,  19856;  Wallin,  1979. 

RECOGNITION.  The  red  colouring  on  the  underside  of  the  abdomen  and  the  conspicuously  pale-tipped  palps 
enable  both  sexes  of  this  species  to  be  distinguished  from  its  relatives.  The  distal  part  of  the  hind  wings  is 
much  more  strongly  smoky  than  in  haemorrhoidalis ,  in  which  the  male  abdomen  is  occasionally  reddish  on 
the  underside. 

In  the  field  the  calling  song  of  the  males  enables  them  to  be  recognized  easily.  Although  quite  similar  in 
basic  pattern  to  those  ofviriduku  and  haemorrhoidalis,  the  song  of  rufipes  differs  in  duration,  lasting  about 
half  as  long  as  that  of  viridulus  and  about  three  times  as  long  as  that  of  haemorrhoidalis. 

SONG  (Figs  34-45) .  The  calling  song  is  an  echeme  usually  lasting  5-10  s  and  consisting  of  syllables 
repeated  at  the  rate  of  about  13-23/s.  The  echeme  begins  quietly  and  gradually  increases  in 
loudness  until  reaching  an  abrupt  end,  thus  resembling  the  first  half  of  the  calling  song  of 
viridulus.  As  in  viridulus  the  syllable  repetition  rate  is  highest  at  the  beginning  of  the  echeme 
(usually  17-23/s),  gradually  lessening  towards  the  end  (when  it  is  usually  13-17/s). 

In  the  presence  of  a  female  the  male  produces  a  courtship  song  quite  similar  to  that  of 
viridulus.  There  is  first  a  series  of  echemes  similar  to  those  of  the  calling  song  but  usually  rather 
longer;  this  is  followed  by  a  quite  different  and  quieter  echeme  lasting  about  5-10  s  and 
composed  of  syllables  repeated  at  the  rate  of  about  10-15/s  (Fig.  37) .  There  are  then  several  loud 
syllables  followed  by  an  attempt  to  copulate  with  the  female.  The  'ticks'  produced  by  backward 
kicks  of  the  hind  tibiae  during  the  courtship  song  of  viridulus  seem  never  to  occur  in  that  of 
rufipes. 

DISTRIBUTION.  This  species  can  tolerate  drier  conditions  than  viridulus  and  often  occurs  in  more  shaded 
habitats.  Its  distribution  is  very  similar,  but  extends  further  south  in  the  Iberian,  Italian  and  Balkan 
peninsulas. 

Omocestus  haemorrhoidalis  (Charpentier) 

(Figs  8,  46-54) 

Gryllus  haemorrhoidalis  Charpentier,  1825:  165.  Syntypes  of  both  sexes,  POLAND/CZECHOSLOVAKIA: 
'Silesia'.  There  are  a  male  (in  bad  condition)  and  two  females  in  the  MNHU,  Berlin  that  have  been 
regarded  as  syntypes  of  this  species,  but  none  is  labelled  'Silesia'  and  there  is  some  doubt  that  they  belong 
to  the  type-series. 

REFERENCES  TO  SONG.  Oscillogram:  Grein,  1984;  Schmidt  &  Schach,  1978.  Diagram:  Bellmann,  1985a; 
Duijm  &  Kruseman,  1983;  Jacobs,  1950, 1953;  Wallin,  1979.  Verbal  description  only:  Faber,  1953;  Harz, 
1957;  Weber,  1984.  Disc  recording:  Grein,  1984.  Cassette  recording:  Bellmann,  1985ft;  Wallin,  1979. 

RECOGNITION.  For  the  distinction  between  this  species  and  petraeus  see  the  remarks  under  that  species. 
Both  sexes  may  be  distinguished  from  viridulus,  rufipes  and  raymondi  by  the  almost  transparent  hind  wings 
(strongly  smoky  in  the  distal  part  in  those  three  species).  In  the  field  the  males  can  be  recognized  quite 
easily  by  their  calling  song,  in  which  the  echemes  are  shorter  than  those  of  viridulus  and  rufipes,  but  much 
longer  than  those  of  raymondi. 

SONG  (Figs  46-54).  The  calling  song  is  an  echeme  lasting  about  2-4  s  and  consisting  of  syllables 


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SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS  233 

repeated  at  the  rate  of  about  25-40/s  (about  double  the  rate  of  viridulus  and  rufipes).  The 
echeme  begins  quietly  but  soon  reaches  maximum  intensity.  The  syllable  repetition  rate 
gradually  lessens  during  the  course  of  the  echeme. 

DISTRIBUTION.  Widely  distributed  in  continental  Europe,  but  not  known  from  the  Scandinavian  mainland 
and  limited  to  mountains  in  the  southern  peninsulas.  The  range  extends  eastwards  across  central  Asia  to 
Mongolia,  Manchuria  and  Korea. 

Omocestus  petraeus  (Brisout) 
(Figs  9,  55-63) 

Acridium  petraeum  Brisout,  1855:  cxiv.  LECTOTYPE  cf ,  FRANCE:  near  Paris,  Lardy  (Brisout)  (IRSNB, 
Brussels),  here  designated  (see  p.  221)  [examined]. 

REFERENCES  TO  SONG.  Diagram:  Luquet,  1978.  Verbal  description  only:  Faber,  1953;  Harz,  1957. 

RECOGNITION.  This  species  lacks  the  smoky  coloration  of  the  distal  part  of  the  hind  wings  shown  by 
viridulus,  rufipes  and  raymondi,  and  is  also  noticeably  smaller.  Distinguishing  it  from  haemorrhoidalis  is 
more  difficult,  especially  if  reliably  identified  specimens  of  both  species  are  not  available  for  comparison. 
O.  petraeus  is  again  noticeably  smaller,  and  the  head  is  larger  in  comparison  to  the  pronotum  and  more 
convex  above  with  shorter  foveolae.  The  males  lack  the  red  colouring  shown  by  the  distal  part  of  the 
abdomen  in  haemorrhoidalis,  showing  a  yellowish  colouring  in  this  region  instead. 

The  highly  distinctive  calling  song,  consisting  of  about  10-20  rapidly  repeated  echemes,  enables  males  of 
this  species  to  be  recognized  easily  in  the  field. 

SONG  (Figs  55-63).  The  calling  song  consists  of  a  sequence  of  about  10-20  echemes  lasting  about 
2-5  s.  The  sequence  begins  quietly,  reaching  maximum  intensity  after  about  6  echemes;  even  at 
its  maximum  intensity  the  song  is  rather  quiet.  The  echeme  repetition  rate  is  usually  about  4/s, 
and  the  syllable  repetition  rate  within  each  echeme  about  80-1 10/s.  Each  of  the  louder  echemes 
lasts  for  about  150  ms  and  contains  about  10-15  syllables. 

In  the  presence  of  a  female  the  male  produces  echemes  of  a  quite  different  kind  (Figs  57,  60, 
63).  Each  echeme  begins  with  a  relatively  long  syllable,  lasting  about  20-50  ms,  and  this  is 
immediately  followed  by  a  series  of  very  short  sounds,  each  lasting  less  than  5  ms,  repeated  at  the 
rate  of  about  50/s.  These  echemes  are  produced  either  singly  or,  more  often,  in  groups  of  about 
2-5. 

DISTRIBUTION.  Western,  central  and  eastern  Europe,  Asia  Minor  and  southern  Siberia.  Largely  absent 
from  the  Iberian  Peninsula,  but  quite  widespread  in  Italy  and  the  Balkan  Peninsula. 

Omocestus  raymondi  ( Yersin) 
(Figs  10,  64-72) 

Stenobothrus  raymondi  Yersin,  1863:  289.  Syntypes  of  both  sexes,  FRANCE:  Var,  near  Hyeres  (Raymond) 
(lost);  the  two  'neotypes'  designated  by  Harz  (1975:  713)  are  clearly  invalid. 

REFERENCES  TO  SONG.  Diagram:  Luquet,  1978.  Verbal  description  only:  Chopard,  1922, 1952. 

RECOGNITION.  This  all-brown  species  is  most  likely  to  be  confused  with  haemorrhoidalis,  but  can  be 
distinguished  from  it  in  both  sexes  by  the  strongly  smoky  distal  part  of  the  hind  wings  (almost  transparent  in 
haemorrhoidalis).  In  the  field  males  may  be  distinguished  from  those  of  haemorrhoidalis  by  the  very  short 
echemes  (lasting  about  1  s)  and  slower  syllable  repetition  rate  (less  than  20/s)  of  the  calling  song. 

SONG  (Figs  64-72).  The  calling  song  is  an  echeme  lasting  about  1-0-1-5  s  and  consisting  of  about 
18-25  syllables  repeated  at  the  rate  of  15-20/s.  Each  echeme  usually  begins  quietly,  rapidly 
increasing  in  intensity.  Oscillograms  show  the  syllables  to  have  a  characteristic  pattern  of  gaps, 
which  often  becomes  obscured  towards  the  end  of  the  echeme  (Figs  67-69).  The  echemes  are 
often  produced  singly  and  repeated  at  irregular  intervals  (10-15  s  is  typical),  but  sometimes  they 
are  in  groups  of  2-4  with  much  shorter  intervals  between  them  (often  2-5  s)  (Figs  65,  66). 

DISTRIBUTION.  Known  only  from  southern  France,  the  Iberian  Peninsula,  north-western  Italy  and  North 
Africa. 


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Omocestus  panteli  (Bolivar) 
(Figs  11, 73-89) 

Stenobothrus  panteli  Bolivar,  1887:  95.  Syntypes  of  both  sexes,  SPAIN:  Cuenca,  near  Ucles  (7MNCN, 
Madrid). 

REFERENCES  TO  SONG.  No  published  work  known  to  me. 

RECOGNITION.  In  the  Iberian  Peninsula,  to  which  it  is  confined,  panteli  is  most  likely  to  be  confused  with 
viridulus;  it  is,  however,  much  smaller  than  that  species  and  can  be  further  distinguished  from  it  by  the  red 
tip  of  the  male  abdomen  and  the  relatively  short  ovipositor.  There  is  also  a  superficial  resemblance  to 
Stenobothrus  stigmaticus  (Rambur),  which  is  similar  in  size  and  also  widespread  in  the  Iberian  Peninsula, 
but  the  pronotal  lateral  carinae  are  straighter  in  panteli,  the  male  cerci  are  conical  throughout  (laterally 
compressed  at  the  tip  in  stigmaticus)  and  the  female  lacks  the  teeth  on  the  ovipositor  shown  by  all  species  of 
Stenobothrus.  There  is  a  large  form  (var.  meridionalis  Bolivar)  found  in  the  south  of  the  peninsula, 
especially  in  mountains;  this  form  is  even  more  like  viridulus,  but  confusion  is  unlikely  as  that  species  does 
not  occur  in  southern  Spain. 

In  the  field  the  song  may  be  recognized  by  its  short  duration  (less  than  2  s)  and  high  syllable  repetition 
rate  (usually  more  than  20/s). 

SONG  (Figs  73-89).  The  calling  song  is  an  echeme  lasting  1-2  s  and  consisting  of  about  30-55 
syllables  repeated  at  the  rate  of  about  18-30/s.  The  echeme  begins  quietly  (sometimes  after  a  few 
louder  syllables)  but  soon  reaches  maximum  intensity.  The  syllable  repetition  rate  gradually 
lessens  during  the  course  of  the  echeme. 

In  the  presence  of  a  female  the  male  first  produces  a  different  kind  of  echeme  lasting  about  2-3 
s,  and  this  is  immediately  followed  by  an  echeme  similar  in  duration  and  syllable  repetition  rate 
to  that  of  the  calling  song  but  with  an  increase  in  intensity  continuing  through  the  whole  echeme 
(Figs  77,  82,  87). 

DISTRIBUTION.  This  species  is  confined  to  the  Iberian  Peninsula,  where  it  is  widespread  and  very  common. 


Omocestus  broelemanni  ( Azam) 
(Figs  90-98) 

Stenobothrus  Brolemanni  Azam,  1906: 128.  LECTOTYPE  cf ,  FRANCE:  Pyrenees-Orientales,  Val  d'Eyne, 
1700-2200  m,  14.viii.1905  (Brolemann)  (MNHN,  Paris),  here  designated  (see  p.  221)  [examined]. 

REFERENCES  TO  SONG.  Oscillogram:  Reynolds,  1986. 

RECOGNITION.  This  Pyrenean  species  is  brachypterous,  the  fore  wings  not  usually  reaching  the  hind  knees  in 
the  male  and  reduced  to  short  lobes  (about  1-5  times  the  length  of  the  pronotum)  in  the  female;  in  both 
sexes  the  hind  wings  fall  short  of  the  fore  wings  by  quite  a  large  gap.  There  is  thus  no  risk  of  confusion  with 
viridulus,  rufipes,  haemorrhoidalis  and  the  other  fully  winged  species  of  Omocestus.  Confusion  is  also 
unlikely  with  the  brachypterous  species  occurring  further  south  in  Spain,  such  as  uhagonii  and  minutissi- 
mus,  which  are  much  smaller  and  not  known  from  the  vicinity  of  the  Pyrenees.  The  relationship  of  O. 
broelemanni  with  O.  antigai  (Bolivar)  and  O.  navasi  Bolivar  has  been  recently  discussed  by  Reynolds 
(1986). 

In  the  field  the  calling  song  of  the  male  is  quite  distinctive,  consisting  of  echemes  lasting  about  as  long  as 
those  of  haemorrhoidalis  but  with  about  half  the  syllable  repetition  rate  of  that  species. 

SONG  (Figs  90-98).  The  calling  song,  recently  described  for  the  first  time  by  Reynolds  (1986),  is  a 
series  of  echemes,  each  lasting  about  1-5-3-0  s  and  consisting  of  about  25-45  syllables  repeated 
at  the  rate  of  about  15-16/s.  Each  echeme  begins  quietly,  reaching  maximum  intensity  after 
about  1  s.  Although  the  echemes  are  sometimes  produced  singly,  they  are  more  often  repeated 
fairly  regularly,  about  every  5-10  s,  in  a  series  of  indefinite  duration. 

During  the  courtship  song,  which  has  also  been  described  by  Reynolds  (1986),  the  male  often 
produces  a  series  of  rather  longer  echemes  (each  lasting  up  to  5  s)  and  this  is  followed  by  a 
variable  number  of  short  echemes  of  small  numbers  of  syllables  (or  even  single  syllables) 


SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS  237 

produced  by  one  leg  only  (Figs  92, 95,  98).  There  is  then  a  series  of  single  syllables  produced  by 
both  legs,  followed  by  an  attempt  at  copulation. 

DISTRIBUTION.  Known  only  from  the  vicinity  of  the  Pyrenees,  particularly  the  eastern  end  of  the  range. 

Omocestus  bolivari  Chopard 
(Figs  99-107) 

Omocestus  Bolivari  Chopard,  1939:  172.  LECTOTYPE  cf ,  SPAIN:  Granada,  Sierra  Nevada,  slopes  of 
Mulhacen,  3000  m  ['2000  m.  environ'  according  to  Chopard,  loc.  cit.],  10.vii.1934  (Balachowsky) 
(MNHN,  Paris),  here  designated  (see  p.  221)  [examined]. 

REFERENCES  TO  SONG.  No  published  work  known  to  me. 

RECOGNITION.  This  species,  known  only  from  the  higher  parts  of  the  Sierra  Nevada  in  southern  Spain,  is 
very  similar  to  uhagonii,  llorenteae  and  minutissimus .  Males  may  be  distinguished  from  all  three  of  these 
species  by  their  short  hind  wings,  which  are  less  than  half  the  length  of  the  fore  wings,  and  the  females  by 
their  short  fore  wings,  which  are  usually  less  than  1-3  times  longer  than  the  pronotum. 

In  the  field  the  isolated  echemes  of  the  calling  song  enable  males  to  be  easily  distinguished  from 
llorenteae  and  minutissimus;  from  uhagonii  they  may  be  distinguished  by  the  much  faster  syllable  repetition 
rate. 

SONG  (Figs  99-107) .  The  calling  song  is  an  echeme  lasting  0-5-2-0  s  and  consisting  of  about  10-30 
syllables  repeated  at  the  rate  of  14— 16/s.  Each  echeme  begins  quietly,  soon  reaching  maximum 
intensity.  Oscillographic  analysis  shows  that  there  are  gaps  in  each  syllable  (mainly  in  the  later 
part  of  the  syllable)  and  that  they  occur  throughout  the  echeme;  there  are,  however,  only  2-4 
gaps  per  syllable  (Figs  102-107),  many  fewer  than  in  uhagonii.  As  in  uhagonii  the  echemes  are 
repeated  at  irregular  intervals,  varying  from  a  few  seconds  to  over  a  minute. 

DISTRIBUTION.  Known  only  from  the  higher  parts  of  the  Sierra  Nevada  in  southern  Spain,  usually  at 
altitudes  above  1500  m. 

Omocestus  uhagonii  (Bolivar) 
(Figs  4, 108-116) 

Gomphocerus  (Stenobothrus)  Uhagonii  Bolivar,  1876:  324.  LECTOTYPE  cf ,  SPAIN:  Madrid,  Navarre- 
donda  (Avila)  (MNCN,  Madrid),  here  designated  (see  p.  221)  [examined]. 

REFERENCES  TO  SONG.  No  published  work  known  to  me. 

RECOGNITION.  Males  of  this  brachypterous  montane  species  can  be  distinguished  from  the  rather  similar 
species  bolivari,  llorenteae  and  minutissimus  by  the  cerci,  which  are  laterally  compressed  towards  the  tip 
(simply  conical  in  the  other  three  species).  Females  can  be  separated  from  bolivari  and  llorenteae  by  the 
length  of  the  fore  wings,  which  are  1-4-1-8  times  longer  than  the  pronotum  (the  corresponding  ratios  for 
bolivari  and  llorenteae  are  0-9-1-3  and  2-1-2-5,  respectively),  and  from  minutissimus  by  the  much  less 
strongly  sigmoid  ventral  profile  of  the  lower  valves  of  the  ovipositor  (cf.  Figs  4,5). 

In  the  field  the  calling  song  of  the  males  enables  this  species  to  be  easily  distinguished  from  llorenteae  and 
minutissimus,  which  produce  echeme-sequences  of  a  quite  different  kind  (cf.  Figs  20-22);  from  bolivari  it 
may  be  distinguished  by  the  much  slower  syllable  repetition  rate  (cf.  Figs  19, 20). 

SONG  (Figs  108-116).  The  calling  song  is  an  echeme  lasting  1-2  s  and  consisting  of  about  10-15 
syllables  repeated  at  the  rate  of  6-7/s.  Each  echeme  begins  quietly,  rapidly  increasing  in 
intensity.  Oscillographic  analysis  shows  that  each  syllable  has  a  large  number  of  gaps  (commonly 
as  many  as  8)  and  that  this  pattern  of  gaps  is  maintained  until  the  end  of  the  echeme  (Figs  111, 
112).  The  echemes  are  repeated  at  irregular  intervals,  varying  from  a  few  seconds  to  over  a 
minute. 

In  the  presence  of  a  female  the  male  produces  a  series  of  slightly  longer  echemes  (lasting  2-3  s) 
at  much  more  regular  intervals  (usually  6-8  in  40  s)  (Fig.  110).  The  echemes  consist  of  about 
15-20  syllables  and  at  the  end  of  some  of  them  (usually  about  half)  there  is  a  group  of  3-4 
syllables  of  a  different  kind  (Figs  113, 116),  repeated  more  rapidly  (at  the  rate  of  about  16/s).  In 


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SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS  243 

the  intervals  between  the  echemes  there  are  quieter  'ticking'  sounds  repeated  fairly  regularly  at 
the  rate  of  about  3-4/s.  The  number  of  echemes  in  the  series  is  variable  but  can  be  more  than  20. 
The  series  of  echemes  is  followed  by  a  variable  period  (often  more  than  a  minute)  in  which  the 
quiet  ticking  continues  and  which  ends  in  a  number  of  sequences  of  a  different  kind  of  sound,  still 
quiet,  and  then  an  attempt  at  copulation.  If  this  is  unsuccessful  the  cycle  begins  again  with 
another  series  of  echemes. 

DISTRIBUTION.  This  species  has  been  previously  known  only  from  the  mountains  of  central  and  northern 
Spain,  but  my  wife  and  I  collected  a  series  of  adults  of  both  sexes  at  El  Chorillo  (2700  m),  south  of  the 
Mulhacen  peak  in  the  Sierra  Nevada,  on  31  July  1983. 

Omocestus  minutissimus  (Bolivar) 
(Figs  5, 117-119) 

Gomphocerus  (Omocestus)  minutissimus  Bolivar,  1878:  424.  Syntypes  of  both  sexes,  SPAIN:  Navarra, 
Cascante  (Perez  Areas);  Madrid,  Escorial  (MNCN,  Madrid;  BMNH,  London;  probably  other  deposi- 
tories) [$  syntype  from  Escorial  in  BMNH  examined]. 

REFERENCES  TO  SONG.  No  published  work  known  to  me. 

RECOGNITION  .  Males  of  this  small  upland  species  may  be  distinguished  from  uhagonii  by  the  cerci ,  which  are 
simply  conical  (laterally  compressed  towards  the  tip  in  uhagonii}.  They  can  be  separated  from  bolivari  and 
llorenteae  by  the  length  of  the  hind  wings,  which  are  more  than  half  the  length  of  the  fore  wings  but  do  not 
reach  the  fore  wing-tips.  Females  can  be  distinguished  from  bolivari  and  llorenteae  by  the  length  of  the  fore 
wings,  which  are  usually  1-4-2-1  times  the  length  of  the  pronotum;  the  corresponding  ratios  for  bolivari  and 
llorenteae  are  0.9-1-3  and  2-1-2-5,  respectively  (these  two  species  are  in  any  case  known  only  from  the 
Sierra  Nevada  or  Sierra  Espuna,  where  minutissimus  does  not  apparently  occur).  Females  of  minutissimus 
may  be  separated  from  those  of  uhagonii  by  the  strongly  sigmoid  ventral  profile  of  the  lower  valves  of  the 
ovipositor  (cf.  Figs  4, 5). 

In  the  field  the  calling  song  of  the  males,  consisting  of  a  rapid  sequence  of  echemes,  enables  this  species 
to  be  easily  distinguished  from  uhagonii  and  bolivari;  from  llorenteae  it  may  be  distinguished  by  being 
shorter  (usually  lasting  less  than  6  s)  and  having  fewer  echemes  in  each  sequence  (usually  fewer  than  10). 

SONG  (Figs  117-119).  The  rather  quiet  calling  song  is  a  sequence  of  about  6-10  echemes  lasting 
about  3-5  s.  The  sequence  begins  very  quietly,  reaching  maximum  intensity  after  about  3-4 
echemes.  The  echemes  are  separated  by  intervals  of  about  100-200  ms,  often  becoming  more 
widely  spaced  towards  the  end  of  the  sequence.  Each  echeme  begins  and  ends  very  quietly, 
giving  a  spindle-shaped  oscillogram  (Fig.  118).  Each  of  the  later  echemes  in  the  sequence  lasts 
about  300-400  ms  and  contains  about  20-30  syllables  repeated  at  the  rate  of  about  80-90/s.  The 
calling  song  is  thus  very  similar  to  that  of  llorenteae,  but  is  quieter  and  shorter,  and  has  fewer 
echemes  composed  of  fewer  syllables. 

DISTRIBUTION.  North-eastern,  central  and  south-eastern  Spain. 

Omocestus  llorenteae  Pascual 
(Figs  123-128) 

Omocestus  llorenteae  Pascual,  1978: 159.  Holotype  O" ,  SPAIN:  Granada,  Sierra  Nevada,  Dornajo,  2000  m, 
19. ix.  1975  (Pascual)  (Universidad  de  Granada)  [examined]. 

REFERENCES  TO  SONG.  No  published  work  known  to  me. 

RECOGNITION.  This  species,  known  only  from  the  Sierra  Nevada  and  Sierra  Espuna  in  southern  Spain,  can 
be  distinguished  from  the  similar  montane  species  uhagonii  and  bolivari  by  the  dark  spots  in  the  medial 
area  of  the  fore  wings  and  by  the  longer  hind  wings,  which  reach  the  tips  of  the  fore  wings;  the  longer  hind 
wings  also  enable  this  species  to  be  distinguished  from  minutissimus,  which  is  in  any  case  not  known  to 
occur  in  the  Sierra  Nevada  or  Sierra  Espuna.  Females  of  llorenteae  also  differ  from  those  of  uhagonii  and 
bolivari  in  that  the  fore  wings  are  more  than  twice  the  length  of  the  pronotum. 

In  the  field  the  calling  song  of  the  males,  consisting  of  a  rapid  sequence  of  echemes,  provides  an  easy 
means  of  distinguishing  this  species  from  most  other  species  of  Omocestus  occurring  in  southern  Spain. 


244 


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SONGS  OF  THE  WESTERN  EUROPEAN  OMOCESTUS 


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From  minutissimus ,  which  has  a  similar  song-pattern,  it  may  be  distinguished  by  being  longer  (usually 
lasting  more  than  6  s)  and  having  more  echemes  in  each  sequence  (usually  more  than  11). 

SONG  (Figs  123-128).  The  calling  song  is  a  sequence  of  about  11-15  echemes  lasting  about  6-8  s. 
The  sequence  begins  quietly,  reaching  maximum  intensity  after  about  5-6  echemes.  The 
echemes  are  usually  separated  by  intervals  of  about  100-200  ms,  often  becoming  more  widely 
spaced  towards  the  end  of  the  sequence.  Each  echeme  begins  and  ends  quietly,  giving  a 
spindle-shaped  oscillogram  (Figs  125,  126).  By  the  middle  of  the  sequence  each  echeme  lasts 
about  350-600  ms  and  contains  about  30-50  syllables  repeated  at  the  rate  of  about  85-95/s.  The 
calling  song  is  thus  remarkably  similar  to  that  of  Myrmeleotettix  maculatus  (Thunberg),  though 
usually  of  shorter  duration  and  composed  of  fewer  echemes. 

DISTRIBUTION.  Known  only  from  the  Sierra  Nevada  and  Sierra  Espuna  in  southern  Spain. 

Omocestus  simony  i  (Krauss) 
(Figs  120-122) 

Stenobothrus  simonyi  Krauss,  1892:  166.  Syntypes  of  both  sexes,  CANARY  ISLANDS:  Lanzarote  (Simony) 
(lost). 

REFERENCES  TO  SONG.  No  published  work  known  to  me. 

RECOGNITION.  This  is  the  only  species  of  Omocestus  known  from  the  Canary  Islands  and  so  in  practice  there 
is  no  problem  in  recognizing  it.  It  is  similar  in  appearance  to  raymondi  but  is  smaller  (total  length  less  than 
14  mm  in  the  male,  less  than  19  mm  in  the  female).  The  calling  song  of  the  male  is  superficially  very  similar 
to  that  of  uhagonii,  oscillographic  analysis  being  required  to  show  that  it  has  far  fewer  gaps  in  each  syllable; 
it  differs  from  that  of  raymondi  in  having  a  much  slower  syllable  repetition  rate. 

SONG  (Figs  120-122).  The  calling  song  is  an  echeme  lasting  about  1-0-2-5  s  and  consisting  of 
about  10-20  syllables  repeated  at  the  rate  of  5-10/s.  Each  echeme  begins  quietly,  reaching 
maximum  intensity  about  midway  through  its  duration.  Oscillographic  analysis  shows  that  there 
are  gaps  in  each  syllable,  similar  in  number  to  those  in  the  calling  song  of  raymondi  (far  fewer 
than  in  uhagonii)  but  not  arranged  in  the  same  pattern  and  maintained  until  the  end  of  the 
echeme  (Figs  121, 122).  The  echemes  are  repeated  at  irregular  intervals. 

DISTRIBUTION.  Known  only  from  the  Canary  Islands,  where  it  occurs  on  Lanzarote  and  Fuerteventura. 

References 

Alexander,  R.  D.  1957.  The  taxonomy  of  the  field  crickets  of  the  eastern  United  States  (Orthoptera: 

Gryllidae:  Acheta).  Annals  of  the  Entomological  Society  of  America  50:  584-602. 

1962.  The  role  of  behavioral  study  in  cricket  classification.  Systematic  Zoology  11:  53-72. 

Alexander,  R.  D.,  Pace,  A.  E.  &  Otte,  D.  1972.  The  singing  insects  of  Michigan.  Great  Lakes  Entomologist 

5:  33-69. 
Alexander,  R.  D.  &  Thomas,  E.  S.  1959.  Systematic  and  behavioral  studies  on  the  crickets  of  the  Nemobius 

fasciatus  group  (Orthoptera:  Gryllidae:  Nemobiinae).  Annals  of  the  Entomological  Society  of  America 

52:  591-605. 
Ander,  K.  1943.  Revision  der  Orthopterensammlungen  Zetterstedts.  Lunds  Universitets  Arsskrift  (N.F., 

Avd.2)38(7),23pp. 

Andrieu,  A.-J.  &  Dumortier,  B.  1963.  Chantes  d'Insectes.  30  cm  disc,  33  rpm.  Neuilly  (Pacific). 
Azam,  J.  1906.  Description  d'un  Stenobothrus  nouveau  des  Pyrenees  [Orth.].  Bulletin  de  la  Societe 

Entomologique  de  France  1906: 128-131. 
Bailey,  W.  J.  1975.  A  review  of  the  African  species  of  Ruspolia  Schulthess  [Orthoptera  Tettigonioidea]. 

Bulletin  de  I'lnstitut  Fondamental  d'Afrique  Noire  (A)  37:  171-226. 
1980.  A  review  of  Australian  Copiphorini  (Orthoptera:  Tettigoniidae:  Conocephalinae).  Australian 

Journal  of  Zoology  27:  1015-1049. 
Bei-Bienko,  G.  Ya.  &  Mishchenko,  L.  L.  1951.  Locusts  and  grasshoppers  of  the  U.S.S.R.  and  adjacent 

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Translations,  Jerusalem,  1964,  iv  +  291  +  xxi  pp.] 


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Beier,  M.  1956.  Feldheuschrecken.  Neue  Brehm-Biicherei  179, 48  pp. 

Bellmann,  H.  1985a.  Heuschrecken.  Beobachten,  Bestimmen.  210  pp.  Melsungen. 

19856.    Die  Stimmen  der  heimischen   Heuschrecken.   Tape   cassette.   Melsungen   (Neumann- 

Neudamm). 
Bolivar,  I.  1876.  Sinopsis  de  los  ortopteros  de  Esparia  y  Portugal.  Segunda  parte.  Anales  de  la  Sociedad 

Espanola  de  Historia  Natural  5:  259-372. 

1878.  Analecta  orthopterologica.  Anales  de  la  Sociedad  Espanola  de  Historia  Natural  7:  423-470. 

1887.  Especies  neuvas  6  criticas  de  ortopteros.  Anales  de  la  Sociedad  Espanola  de  Historia  Natural  16: 

90-114. 

1897.  Catalogo  sinoptico  de  los  ortopteros  de  la  fauna  iberica  [second  part].  Annaes  de  Sciencias 


Natvraes  4;  2X8-232. 

Borck,  J.  B.  von  1848.  Skandinaviens  ratvingade  insekters  natural-historia.  xvi  +  146  pp.  Lund. 
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France  (3)  3:  cxiv-cxvi. 
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sur  disque  reproduit  a  des  vitesses  ralenties  [pp.  82-88].  In  Busnel,  R.-G.  [Ed.],  Colloque  sur 

1'acoustique  des  Orthopteres.  Annales  des  Epiphyties  (fascicule  special  de  1954),  448  pp. 
Busnel,  R.-G.  1955.  Sur  certains  rapports  entre  le  moyen  d'information  acoustique  et  le  comportement 

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Orthopteres.  Annales  des  Epiphyties  (fascicule  special  de  1954),  448  pp. 

Charpentier,  T.  de  1825.  Horae  entomologicae,  adjectis  tabulis  novem  coloratis.  xvi  +  262  pp.  Bratislava. 
Chopard,  L.  1922.  Orthopteres  et  Dermapteres.  Faune  de  France  3,  vi  +  212  pp. 
1939.  Description  d'une  espece  nouvelle  du  genre  Omocestus  [Orth.  Acrididae].  Bulletin  de  la  Societe 

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249 


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Invalid  names  are  in  italics; 
principal  page  references  in 
bold. 

Acheta  220 
alluaudi  222 
antigai  214, 222,  236 
aymonissabaudiae  222 

biguttulus  214, 215, 220 
bolivari  214, 220,  221,  222, 226, 

237, 243 

broelemanni  221, 222,  224,  236 
brunneus  214,  215 
burri  214, 222 

caucasicus  222 

Chorthippus  214,  215, 220, 221 
corsicus  214, 222 
cuonaensis  222 

demokidovi  222 
Dirshius  221 

enitor  222 
Euchorthippus  220 

femoralis  214,  222 
Gomphocerippus  221 


INDEX 

Gomphocerus  221 
grossum  226 

haemorrhoidalis  213,  222,  224, 

226,  227,  233,  236 
heymonsi  222 
hingstoni  222 

kaestneri  214,  222 
knipperi  214,  222 

lecerfi  222 

lepineyi  222 

llorenteae  214,  220, 222, 223,  224, 

237, 243 

lopadusae  214, 222 
lucasii  222 

maculatus  246 
megaoculus  222 
meridionalis  236 
minutissimus  214, 220,  222, 223, 

224,  236,  237, 243,  246 
minutus  222 
mollis  215 
motuoensis  222 
Myrmeleotettix  221,  246 

nanus  222 

navasi  214,  222, 236 

Nemobius  220 


Neoconocephalus  220 
nyalamus  222 

panteli  214, 219, 222, 223,  226,  236 
petraeus  213, 214, 220,  221, 222, 

223,224,227,233 
Poecilimon  220 
pullus221 

raymondi  214, 222, 224, 226, 227, 

233, 246 
rufipes  213, 219,  220, 222,  224, 

226,  227, 233,  236 

simonyi  214,  222, 224,  246 
Stenobothrus221,236 
Stethophyma  226 
stigmaticus  236 

tibetanus  222 
tzendsureni  222 


uhagonii  214, 220,  221, 222, 223, 

224, 236, 237,  243,  246 
uvarovi214,221,222 

ventralis  227 

viridulus  214,  219, 220,  221,  222, 
224, 226,  227, 233, 236 

znojkoi  222 


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The  songs  of  the  western  European  grasshoppers  of  the  genus  Omocestus  in  relation  to  their 
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19 

PRE 


Bulletin  of  the 

British  Museum  (Natural  History) 


The  structure  and  affinities  of  the 
Hedyloidea:  a  new  concept  of  the 
butterflies 

M.  J.  Scoble 


Entomology  series 

Vol53  No  5  18  December  1986 


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ISBN  0  565  06023  6 

ISSN  0524-6431  Entomology  series 

Vol53No5pp251-286 
British  Museum  (Natural  History) 
Cromwell  Road 
London  SW7  5BD  Issued  18  December  1986 


The  structure  and  affinities  of  the  Hedyloidea:  a 

new  concept  of  the  butterflies 

.. 

M.  J.  Scoble 

Department  of  Entomology,  British  Museum  (Natural  History),  Cromwell  Road,  London  SW7 
5BD 


Contents 

Synopsis  ....................................................................................................  25  1 

Introduction  ..............................................................................................  25  1 

Material  .................................................................................................  253 

Hedylidae  Guenee  ......................................................................................  253 

Generic  synonymy  ....................................................................................  254 

Macrosoma  Hiibner  .....................................................................................  254 

General  appearance  .....................................................................................  254 

Description  ................................................................................................  254 

Comments  on  selected  structures  ....................................................................  260 

Adult  ....................................................................................................  260 

Juvenile  stages  .........................................................................................  280 

Discussion  .................................................................................................  281 

Epilogue  :  a  suggested  phylogeny  .....................................................................  284 

Acknowledgements  .....................................................................................  284 

References  ................................................................................................  284 

Synopsis 

The  affinities  of  the  Hedylidae,  a  group  of  about  40  Central  and  South  American  Lepidoptera,  are 
reassessed.  Currently,  the  taxon  is  treated  as  a  tribe  ('Hedylicae')  of  the  Oenochrominae  (Geometridae), 
but  evidence  is  presented  here  to  show  that  hedylids  are  butterflies  although  they  look  moth-like  in  many 
respects.  The  five  generic  names  in  the  family  are  synonymized  in  this  work  with  the  oldest,  Macrosoma 
Hiibner. 

The  taxonomic  history  of  the  Hedylidae  is  discussed,  and  the  reasons  for  the  proposed  assignment  of  the 
family  to  the  Rhopalocera  are  given.  A  description  of  the  family  is  followed  by  comments  on  characters 
selected  for  their  biological  or  systematic  interest.  In  this  section  comparisons  are  made  between  the 
Hedylidae  and  other  Lepidoptera,  particularly  the  butterflies,  and  their  phylogenetic  relationships  are 
discussed.  Hedylidae  +  Hesperioidea  -I-  Papilionoidea  (s.str.)  probably  form  a  monophyletic  taxon, 
although  doubts  have  been  expressed  about  the  relationship  between  skippers  and  true  butterflies.  It  is 
tentatively  suggested  that  within  the  Rhopalocera,  the  Hedylidae  are  more  closely  related  to  the 
Papilionoidea  than  are  the  Hesperiidae  to  the  Papilionoidea;  that  is,  the  butterflies  without  the  Hedylidae 
are  paraphyletic. 

The  evidence  of  the  butterfly  affinities  of  hedylids  comes  from  adults,  larvae,  and  pupae;  the  structure  of 
the  thorax  and  the  base  of  the  abdomen  of  the  adult  was  found  to  be  particularly  revealing. 

Key  words:  Lepidoptera,  Rhopalocera,  Hedyloidea  stat.  n.,  Hedylidae,  Geometridae,  morphology, 
phylogeny. 

Introduction 

Guenee  (1857)  described  the  Hedylidae  as  one  of  the  26  component  families  of  the  Phalenites. 
The  Phalenites  are,  with  modification,  the  equivalent  of  what  are  now  called  the  Geometridae. 
The  Hedylidae  were  treated  by  Prout  (1910;  1931)  as  a  tribe  ('Hedylicae')  of  the  Oenochromi- 
nae (Geometridae),  and  no  reassessment  of  the  relationships  of  the  group  has  been  attempted 
subsequently.  In  fact  the  Hedylidae  are  misplaced  in  the  Geometridae,  the  most  obvious  reason 
being  the  absence  of  tympanal  organs  from  the  base  of  the  abdomen  in  the  former  -  structures 
that  are  characteristic  of  the  latter. 


Bull.  Br.  Mus.  nat.  Hist.  (Ent.)  53  (5):  251-286  Issued  18  December  1986 


252  M.  J.  SCOBLE 

The  Hedylidae  consist  of  some  40  species,  from  South  or  Central  America.  Below,  I  argue 
that  not  only  is  this  group  misplaced  in  the  Geometridae  but  also  that  its  phylogenetic  affinities 
are  with  the  butterflies  (Rhopalocera:  Hesperioidea  or  skippers,  and  Papilionoidea  or  true 
butterflies)  despite  the  moth-like  external  appearance  of  the  adult  insects. 

Both  Guenee  (1857)  and  Prout  (1910;  1931)  noted  certain  similarities  between  hedylids  and 
butterflies.  Guenee  even  went  so  far  as  to  name  one  species  napiaria  because  it  resembled  the 
pierid  butterfly  Pieris  napi  (L.).  He  named  another  species  heliconiaria  since  it  reminded  him  of 
certain  species  of  the  papilionoid  genus  Heliconius  Kluk.  Prout  (1910;  1931)  noted  the 
butterfly-like  shape  of  the  adults,  and  the  remarkable  girdled  pupa  in  the  two  species  for  which 
he  had  information.  However,  neither  of  these  workers  suggested  that  these  features  meant  that 
hedylids  and  butterflies  were  related,  and  they  presumably  regarded  the  similarities  as  indepen- 
dent derivations.  Prout  (1931)  was  clearly  uncomfortable  about  the  geometrid  affinities  of  the 
group  since  he  suggested  that  the  'Hedylicae'  might  deserve  even  a  separate  family  status. 
However,  neither  he  nor  Guenee  apparently  considered  that  the  butterfly-like  features  they 
observed  might  actually  be  homologous  in  the  taxa  concerned  and  thus  indicative  of  true  affinity. 
Why? 

There  are  two  reasons,  one  philosophical  and  one  practical.  We  tend  to  develop  a  conception 
of  a  given  taxon.  We  think  of  butterflies  as  a  package,  and  develop  what  might  be  termed  a 
butterfly  Gestalt  -  colourful,  diurnal  lepidopterans  with  clubbed  antennae  and  amplexiform 
rather  than  frenate  wing-coupling.  This  is  not  to  say  that  we  do  not  recognize  exceptions.  Males 
of  the  skipper  Euschemon  Doubleday  have  a  frenulum  and  retinaculum;  satyrines  generally  are 
not  brightly  coloured,  many  butterflies  do  not  have  strongly  clubbed  antennae,  and  a  few  are 
nocturnal.  By  definition,  exceptions  do  not  destroy  a  Gestalt.  But  hedylids  do  not  conform  to  the 
traditional  butterfly  package.  They  do  not  have  clubbed  antennae;  in  some  species  these 
structures  are  even  bipectinate.  In  males  of  most  species  the  frenulum  and  retinaculum  are  very 
well  developed,  and  the  wing  pattern  in  most  species  bears  little  resemblance  to  that  of 
butterflies  (Figs  1-18).  Most  species  seem  to  be  nocturnal. 

Most  taxa  are  diagnosed  by  a  set  of  attributes  rather  than  by  a  single  character  or  very  few 
characters.  Even  so,  such  diagnoses  often  need  qualification  to  incorporate  those  taxa  that  do 
not  fit  -  for  example,  primitive  species  that  have  not  yet  acquired  all  the  attributes  of  the 
majority,  or  those  specialized  species  that  have  lost  them  secondarily.  Crowson  (1970),  noting 
the  essentialist  nature  of  such  diagnoses,  called  the  groups  Aristotelian  to  distinguish  them  from 
taxa  delimited  phylogenetically.  Crowson  also  coined  the  phrase  'the  non-congruence  principle' 
to  suggest  that  it  is  most  unlikely  that  a  taxon  can  be  defined  on  more  than  one  or  two  characters, 
since  when  a  taxon  originates  many  of  the  characters  found  in  later  members  of  the  group  have 
not  yet  evolved.  Characters  we  regard  as  typical  of  a  given  taxon  do  not  evolve  at  once. 
Therefore,  in  the  present  case,  we  should  not  expect  the  ground  plan  of  the  butterflies  as  a  whole 
or  even  the  true  butterflies  to  exhibit  the  characteristics  of  the  more  advanced  members  of  the 
group.  It  should  not,  then,  be  unexpected  to  find  a  group  strongly  affiliated  to  the  butterflies  that 
does  not  fit  our  conception  of  them .  However ,  if  it  is  our  stated  aim  to  produce ,  as  far  as  possible , 
a  phylogenetic  classification,  then  merely  one  derived  character  shared  between  two  taxa 
provides  evidence  of  monophyly.  In  fact,  the  Hedylidae  possess  several  such  characters. 

Having  rejected  the  idea  that  hedylids  might  be  related  to  butterflies,  neither  Guenee  nor 
Prout  looked  further.  This  may  largely  have  been  because  morphological  studies  were  less 
extensive  in  Guenee's  time  than  they  are  today.  Much  the  same  can  be  said  of  Prout,  who  had  a 
remarkable  ability  to  observe  structures  and  their  significance  on  dried  specimens,  but  did  little 
dissection.  (Only  late  in  his  career  did  he  study  genitalia.)  In  fact  many  of  the  characters  that 
support  the  suggestion  that  hedylids  are  butterflies  are  to  be  observed  only  after  detailed 
examination  of  the  sclerites  of  the  thorax  and  the  base  of  the  abdomen. 

The  present  study  evolved  from  a  project  to  re-diagnose  the  Oenochrominae  (Geometridae), 
and  to  exclude  those  groups  that  currently  make  the  subfamily  unnatural  (non-monophyletic). 
The  extraordinary  combination  of  attributes  of  the  Hedylidae  demonstrates  that  their  affinities 
lie  neither  with  Oenochroma  Guenee  (the  type  genus  of  the  Oenochrominae)  and  its  relatives, 
nor  with  the  Geometridae  as  a  whole.  In  particular,  abdominal  tympanal  organs,  which  are 


NEW  CONCEPT  OF  BUTTERFLIES  253 

typical  of  geometrids,  are  absent  from  hedylids.  Also,  prolegs  are  present  on  abdominal 
segments  3  to  6  and  10  of  the  larva,  unlike  most  geometrids  where  their  number  is  reduced.  The 
first  tergum  of  the  abdomen  is  strongly  'pouched'  (a  condition  described  for  papilionoids  by 
Ehrlich,  1958&),  and  pre-  and  postspiracular  bars  are  present.  The  abdomen  is  curved  and 
flattened  laterally,  particularly  in  the  male.  The  remarkable  pupa  is  attached  to  a  leaf  by  a  silken 
girdle:  there  is  no  cocoon.  The  apical  tibial  spurs  of  the  hindleg  are  absent  in  all  species  except 
the  male  of  one.  The  precoxal  (or  paracoxal)  sulcus  of  the  mesothorax  joins  the  'marginopleural' 
sulcus.  In  the  metathorax  the  furcal  apophysis  is  weakly  sagittate  in  dorsal  view.  The  pretarsus  is 
lost  from  the  forelegs  of  the  male  although  minute  claws  are  present,  and  the  tarsomeres  are 
reduced  to  two  by  fusion.  The  head  of  the  larva  is  extended  into  a  pair  of  extremely  long  horns, 
and  there  is  an  anal  comb  on  abdominal  segment  10.  These  characters  are  all  represented 
in  various  butterflies.  In  contrast,  the  moth-like  features  include  the  absence  of  clubs  on 
the  antennae,  and  a  well-developed  frenulum  and  retinaculum  in  the  males  of  most 
species,  although  they  are  reduced  and  functionless  in  a  few.  The  wing  venation  is  peculiar: 
Rsi  and  Rs4  share  a  common  stem,  but  Rs2  is  separate  from  the  cell  and  is  not  stalked  with  Rs3. 
Rsi  and  Rs^  are  distinctly  sinuate.  Special  modifications  of  the  wings,  including  a  complex  of 
small  chambers  at  the  base  of  the  forewing  and  a  prominent  flap  extending  from  Sc,  are 
described  below. 

The  body  of  this  work  aims  to  provide  the  evidence  that  suggests  that  hedylids  are  butterflies. 
Are  the  apparent  butterfly  attributes  genuinely  shared-derived  characters  of  hedylids  and 
butterflies  or  are  they  convergent  or  parallel  developments?  In  short,  are  the  Hedylidae  more 
closely  related  to  the  butterflies  than  to  any  other  lepidopterans? 

A  further  question  is:  are  the  Hedylidae  more  closely  related  to  the  true  butterflies 
(Papilionoidea)  than  to  the  skippers  (Hesperioidea),  or  does  the  family  represent  the  sister 
group  of  the  Rhopalocera  as  a  whole? 

A  general  description  of  the  characters  found  in  this  family  is  provided,  together  with  a 
summary  of  its  biology  and  distribution.  This  is  followed  by  observations  on  selected  structures 
with  comments  on  their  systematic  importance.  Finally,  the  evidence  for  the  affinity  of  the 
Hedylidae  with  the  Hesperioidea  and  the  Papilionoidea  is  assessed. 

Material 

The  observations  were  based  on  specimens  housed  in  the  British  Museum  (Natural  History). 
Juvenile  stages  of  one  species  were  lent  by  Mr  R.  O.  Kendall,  San  Antonio,  Texas. 

Examination  of  the  structure  of  the  thorax  was  necessarily  limited  to  avoid  breaking  up 
numerous  specimens.  From  the  study  of  more  easily  observed  characters  the  family  Hedylidae 
appears  to  be  a  homogeneous  group,  and  thoracic  structures  probably  do  not  differ  significantly 
between  species.  Thoracic  structure  was  examined  in  species  that  showed  most  structural 
variation  in  the  genitalia. 

HEDYLIDAE  Guenee 

Hedylidae  Guenee,  1857:  521.  Type  genus:  Hedyle  Guenee,  1857:  521. 

Guenee  (1857)  described  the  family  for  three  genera,  Hedyle  Guenee,  Venodes  Guenee,  and 
Phellinodes  Guenee.  With  the  exception  of  Prout  (1910;  1931),  most  authors  have  been  content 
to  describe  new  species  or  genera  without  discussing  the  taxonomic  relationships  of  the  group  as 
a  whole.  Prout,  in  both  these  works,  dealt  with  all  the  genera  and  species.  He  treated  the  taxon 
as  a  tribe  -  the  'Hedylicae'  -  of  the  subfamily  Oenochrominae  (Geometridae).  Authors  who 
preceded  Prout  assigned  the  genera  to  various  infrafamilial  categories  of  the  Geometridae, 
some  of  which  are  not  currently  in  use,  with  no  justification  for  their  action.  Kendall  (1976),  in 
his  notes  on  the  life-history  of  one  species,  used  the  name  Hedylidae,  therefore  effectively 
reviving  Guenee's  original  status. 

The  use  of  the  name  Hedylidae  is  fully  justified  by  Guenee's  priority,  Kendall's  subsequent 
usage,  and  the  absence  of  any  competing  name. 


254  M.  J.  SCOBLE 

Generic  synonymy 

The  species  of  the  family  show  no  fundamental  structural  difference  from  each  other,  and  form  a 
highly  compact  group,  particularly  in  the  male  genitalia,  wing  venation,  wing  shape,  legs  and 
antennae.  Below,  I  synonymize  the  five  genera  that  are  currently  accepted.  Three  of  these  are 
monotypic. 

Macrosoma  tipulata  Hiibner  and  Lasiopates  hyacinthina  Warren  are  species  that  exhibit  the 
greatest  differences  from  the  majority  of  the  family.  The  valvae  of  the  male  genitalia  of 
hyacinthina  (Fig.  79)  are  of  a  more  complex  shape,  the  two  components  of  the  gnathos  are  not 
fused  medially,  and  the  juxta  is  characteristic. 

One  reason  for  the  proliferation  of  genera  in  the  Lepidoptera  is  that  since  the  study  of  the 
male  genitalia  became  universal,  there  has  been  a  strong  tendency  to  treat  as  genera  those 
groups  where  there  is  a  morphological  'gap'  in  these  structures.  The  consequence  has  been  that 
although  a  family  may  contain  a  few  large  genera  there  are  usually  several  anomalous  species.  If 
these  are  then  excluded  from  one  of  the  main  genera  each  has  to  be  assigned  to  a  new  genus.  In  a 
family  with  several  large  genera  it  may  be  impossible  to  decide  to  which  genus  such  'outlier' 
species  should  be  assigned.  The  description  of  a  new  genus  therefore  becomes  inevitable.  In  a 
family  with  one  large  genus  there  is  often  a  good  case  to  be  made  for  including  all  species  within 
that  genus  and  using  lower  categorical  rank,  e.g.  subgenus,  or  simply  species-group,  to 
distinguish  outliers.  In  the  present  case,  despite  the  synonymy,  I  have  not  so  divided  Macrosoma 
in  this  way.  This  is  best  left  until  such  time  as  a  revision  of  all  the  species  is  undertaken.  The 
synonymized  names  can  then  be  used  as  subgenera,  if  required. 

MACROSOMA  Hiibner 

Epirrita  Hiibner,  1808.  [Name  unavailable,  see  Fletcher,  1979.] 

Macrosoma  Hiibner,  1818:  10.  Type  species:  Macrosoma  tipulata  Hiibner,  1818: 10  (by  monotypy). 

Hedyle  Guenee,  1857:  521.  Type  species:  Hedyle  heliconiaria  Guenee,  1857  (by  monotypy).  [Four 
described  species.]  Syn.  n. 

Venodes  Guenee,  1857:  522.  Type  species:  Venodes  napiaria  Guenee,  1857:  522  (by  monotypy).  [Mono- 
typic.] Syn.  n. 

Phellinodes  Guenee,  1857:  523.  Type  species:  Phellinodes  satellitiata  Guenee,  1857:  523  (by  monotypy). 
[Thirty-three  described  species.]  Syn.  n. 

Macrophila  Walker,  1862:  1463,  1465.  Type  species:  Macrosoma  tipulata  Hiibner,  1818  (by  monotypy). 
[Junior  objective  synonym  of  Macrosoma,  see  Fletcher,  1979.] 

Hyphedyle  Warren,  1894:  375.  Type  species:  Hedyle  rubedinaria  Walker,  1862:  1464  (by  original 
designation).  [Synonymized  with  Hedyle  Guenee  by  Prout,  1910.] 

Lasiopates  Warren,  1905:  310.  Type  species:  Lasiopates  hyacinthina  Warren,  1905:  311  (by  original 
designation).  [Monotypic.]  Syn.  n. 

Hedylidae  remains  the  valid  name  of  the  taxon  since  family  names  are  not  rejected  when,  after  1960, 
their  type  genera  are  found  to  be  junior  synonyms. 

General  appearance 

Hedylids  are  fairly  delicate  insects  with  curved  abdomens,  particularly  in  the  male  (Fig.  63).  Figs 
1-18  show  the  main  variation  in  wing  patterns:  many  of  the  species  bear  semi-transparent 
patches  on  the  wings.  The  fore  wings  are  weakly  or  strongly  emarginate  at  the  apex.  The  larva 
(Figs  19,  100,  101)  bears  a  pair  of  very  long  'horns'  on  the  head  capsule,  prolegs  on  abdominal 
segments  3  to  6  and  10,  a  pair  of  furcae  on  the  anal  segment,  and  an  anal  comb.  The  pupa  (Figs 
20,  21)  lacks  a  cocoon,  and  is  attached  to  its  substrate  by  a  silken  girdle. 

Description 

Adult 

Head  (Figs  22-29).  Frons  narrow  and  protuberant  (Figs  22,  23).  Compound  eyes  large.  Ocelli  absent. 
Chaetosemata  present,  one  chaetosema  behind  each  eye  (Fig.  22).  Antenna  (Figs  25-28):  bases  close 


NEW  CONCEPT  OF  BUTTERFLIES 


255 


1 


Figs  1-6  Adults  of  Hedylidae.  1,  Macrosoma  heliconiaria  (Guenee),  x  1-5;  2,  M.  hyacinthina  (Warren), 
x  1-1;  3,  M.  tipulata  Hiibner,  x  1-1;  4,  M.  rubedinaria  (Walker),  x  1-5;  5,  M.  ustrinaria  (Herrich- 
Schaffer),  x  1-5;  6,  M.  leucophasiata  (Thierry  Mieg),  x  1-5. 


together;  filiform  (Figs  26,  27)  or  bipectinate  (Fig.  25);  scaled  dorsally.  Pilifers  present.  Maxillae:  palpi 
minute,  one-segmented;  galeae  form  a  well-developed  proboscis,  not  scaled  at  base.  Labial  palpi 
ascending  (Fig.  24),  three-segmented  with  a  deep  sensory  invagination  on  distal  segment  (Fig.  29). 
Tentorium:  medial  swelling  on  each  tentorial  arm  (Fig.  23),  but  dorsal  arms  not  developed. 

Thorax  (Figs  30,  31).  Prothorax  with  membranous  patagia  and  parapatagia;  patagia  protuberant. 
Mesothorax:  anepisternum  well  developed,  not  reduced  (Fig.  30);  precoxal  sulcus  (paracoxal  sulcus  of 
Scott,  1985)  joins  'marginopleural'  suture  (Fig.  31).  Metathorax:  furcal  apophyses  weakly  sagittate  in 
dorsal  view;  dorsal  lamellae  of  secondary  furcal  arms  fused  mesally  in  their  extension  anteriorly  from  the 
arms  to  the  furcal  apophyses. 

Wing  venation  (Fig.  32).  (Following  the  recommendations  of  Wootton  (1979),  in  this  work  I  label  the 


256 


M.  J.  SCOBLE 


8 


10 


11 


12 


Figs  7-12  Adults  of  Hedylidae.  7,  Macrosoma  subornata  (Warren),  x  1-5;  8,  M.  paularia  (Schaus),  X 
1-5;  9,  M.  coscoja  (Dognin),  x  1-5;  10,  M.  satellitiata satellitiata  (Guenee),  x  1  •  1 ,  hyaline  patch  arrowed; 
11,  M.  albifascia  albifascia  (Warren)  x  1-1;  12,  M.  hedylaria  (Warren),  x  1-1. 


branches  of  the  radial  sector  as  Rsi,  Rs2,  etc.,  to  distinguish  them  from  the  radius  (/?i).)  Fore  wing: 
accessory  cells  absent.  Rs\  and  Rs2  sinuous  after  branching  from  cell;  Rs4  stalked  with  /?s3,  Rs2  not  stalked 
with  Rs3  (although  it  shares  a  common  stem  with  Rs3  and  Rs4  before  the  three  branches  arise);  Rs4  remote 
from  MI;  CuP  often  weakly  indicated;  M2  about  equal  distance  from  Ml  as  from  M2;  anals  forked  at  base. 
Hind  wing:  Rs  separate  from  Sc+R]  from  near  base  of  wing;  M2  about  equidistant  from  MI  and  M3. 

Wing  coupling  (Figs  33-37).  Male  usually  with  strong  frenulum  and  a  very  long,  well-developed 
retinaculum;  frenulum  occasionally  reduced  and  non-functional  (Fig.  35).  Female:  frenulum  composed  of 
a  few  bristles  (Fig.  36);  retinaculum  absent. 

Wings  .-pattern  and  specializations.  Figs  1-18  illustrate  the  variety  of  shape  and  pattern.  Vestiture:  rather 
weakly  scaled;  prominent  piliform  scales  present  particularly  near  base  of  wings  (Fig.  38),  and  more 


NEW  CONCEPT  OF  BUTTERFLIES 


257 


13 


14 


15 


16 


18 

Figs  13-18  Adults  of  Hedylidae.  13,  Macrosoma  lucivittata  (Walker),  cf,  x  1-1;  14,  M.  lucivittata 
(Walker),  $,  x  1-1;  15, M.  nigrimacula(  Warren),  x  1-1;  16, M.  leptosiata  (Felder) ,  x  1-1,  hyaline  patch 
arrowed;  17,  M.  napiaria  (Guenee),  x  1-1;  18,  M.  hedylaria  (Warren),  ventral  surface  with  wings  folded 
vertically,  x  1-5. 

numerous  on  hind  wing  than  fore  wing.  Upperside  and  underside  with  the  same  pattern.  Areas  of 
semi-transparent  scales  often  present.  Fore  wing  long,  not  broad,  weakly  falcate  often  with  apex  weakly 
emarginate;  posterior  edge  expanded  into  a  small  lobe  at  base  of  wing  in  males  of  some  species  (Fig.  43), 
sometimes  merely  a  very  weak  expansion.  Holding  area  (Haftfeld)  not  present  on  ventral  surface  (nor  on 
metascutum).  Two  small,  but  prominent  chambers  at  base  of  wing  (Figs  37,  42);  one,  predominantly  in  the 


258 


M.  J.  SCOBLE 


19 


20 


21 


Figs  19-21     Larva  and  pupa  of  Hedylidae.  19,  larva  of  Macrosoma  heliconiaria  (Guenee),  x  3-5;  20,  21. 
pupal  exuviae  of  (20)  M.  lucivittata  (Walker),  x  1-5;  (21)  M.  nigrimacula  (Warren),  x  1-5. 


ventral  plane,  is  a  distension  of  base  of  Sc,  the  other,  predominantly  in  the  dorsal  plane,  a  distension  of  Cu. 
Prominent  ventral  fold  expands  from  Sc  on  underside  (Figs  39,  40).  Scale-enclosed  pocket  present  at  very 
base  of  underside  of  wing  (Figs  40,  41).  Hindwing  rounded:  in  males  often  with  folded  area  (of  glassy 
appearance)  present  (Figs  10,  16,  44,  47);  basally  usually  with  associated  sclerotized  protuberances  (Figs 
44-47, 50)  and  specialized  scales  (Figs  48, 49);  base  of  costa  weakly  or,  when  frenulum  is  reduced,  strongly 
expanded  into  a  'shoulder'  (Fig.  35),  composed  of  extended  frenulum-plate  and  costal  region. 

Base  of  fore  wing  (Figs  37,  51).  Second  median  plate  with  substantial  part  hidden  under  base  of  anal 
veins. 

Legs  (Figs  52-62).  Microtrichia  present  on  ventral  surface  at  base  of  tarsi  of  mid  leg  and  hind  leg  of  male 
and  all  legs  of  female. 

Fore  leg.  Epiphysis  present  (Figs  52-54);  tibial  spurs  absent.  Male:  tarsus  (Figs  52,  55,  57,  58) 
two-segmented,  number  reduced  by  fusion  but  tarsus  not  shortened;  pretarsus  reduced  to  pair  of  minute 
claws  (Fig.  58),  empodium  and  pulvilli  absent;  tarsi  not  spined,  but  hairy;  occasionally  with  a  scent  brush. 
Female:  tarsomeres  not  fused,  tarsi  five-segmented  (Fig.  54);  pretarsus  not  reduced,  bears  a  pair  of 
strongly  curved  claws  (Fig.  59)  that  are  weakly  and  asymmetrically  forked;  pulvilli  present;  tarsus  spined 
ventrally,  not  hairy. 

Mid  leg:  with  one  pair  of  tibial  spurs;  tarsus  five-segmented,  tarsomeres  not  fused;  pretarsus  not 
reduced,  tarsal  claws  weakly  forked;  tarsus  spined  on  ventral  surface. 

Hind  leg:  with  one  pair  (Fig.  61)  or  occasionally  two  pairs  (Fig.  62)  of  tibial  spurs;  tibia  somewhat 
swollen;  otherwise  as  mid  leg. 

Abdomen.  Weakly  to  moderately  curved  (Fig.  63),  flattened  laterally  not  dorso-ventrally,  and  narrow- 
ing slightly  towards  base.  Tympanal  organs  absent. 

Pregenital  abdomen  (Figs  64,  65, 68).  Segment  I  with  narrow  prespiracular  bar'and  broad  postspiracular 
bar  (tergopleural).  Tergum  I  with  large  membranous  pouch  and  strong  tergal  braces.  Sternum  II  with  pair 
of  mediolateral  sclerotizations  anteriorly;  sternal  apodemes  reduced. 


NEW  CONCEPT  OF  BUTTERFLIES  259 

Male  postabdomen  (Figs  66,  67).  Tergum  VIII  longer  than  sternum  VIII  forming  a  short  hood. 

Female  postabdomen .  Segment  VII  in  form  of  a  short  hood  over  the  genitalia ;  tergum  VII  large ;  sternum 
VII  shorter,  usually  bearing  a  series  of  prominent  setae  on  fold  just  anterior  to  the  ostium  bursae  (Fig.  90). 

Male  genitalia  (Figs  69-80).  Genital  capsule  is  'deep'  dorso-ventrally  owing  to  length  of  lateral  arms  of 
vinculum.  Segment  IX  comprises  a  completely  sclerotized  ring,  i.e.  tergum  and  sternum  fused.  Tegumen 
narrow,  sometimes  prominently  bilobed.  Uncus  large,  usually  triangular  in  ventral  view  since  it  narrows 
apically;  alternatively  apex  is  truncated  or  otherwise  modified  (Fig.  83).  Vinculum:  ventral  plate  narrow 
and  produced  into  a  prominent  narrow  saccus;  rarely,  saccus  is  rounded.  Gnathos  usually  large  and  heavily 
sclerotized;  occasionally  lateral  arms  do  not  meet  medially.  Valvae  (Figs  77-82)  simple,  undivided,  usually 
approximately  triangular.  Diaphragm  unsclerotized,  or  occasionally  sclerotized  as  a  juxta  into  a  small 
simple  plate  or,  rarely,  a  prominent  U-shaped  structure  (Fig.  84).  Aedeagus  (Figs  85-89)  a  simple  tube; 
vesica  unmarked,  or  with  weak  sclerotizations. 

Female  genitalia  (Figs  90-99).  Tergum  VIII  a  well-sclerotized  band.  Anal  papillae  usually  in  form  of  a 
pair  of  ear-like  lobes.  Anterior  apophyses  sometimes  reduced  (Figs  93,  96).  Posterior  apophyses 
developed  normally.  Bursa  copulatrix:  a  narrow  ductus  leads  into  a  globose  or  sub-globose  corpus;  signum 
absent  or  present;  if  present  then  of  a  characteristic  shape  (Figs  98, 99). 

Egg 

An  egg,  found  while  dissecting  the  female  genitalia  of  a  specimen,  was  oval. 

Larva 

Based  on  M.  heliconiaria  (comb,  n.)  (Fig.  19,  and  see  Kendall,  1976),  with  further  information  on  M. 
nigrimacula  (Warren)  (comb,  n.)  derived  from  larval  exuviae  housed  in  the  British  Museum  (Natural 
History).  When  this  study  was  at  an  advanced  stage,  Mr  T.  Fox  (London  Butterfly  House)  provided  a 
colour  transparency  of  a  larva  (subsequently  reared  through)  of  M.  cascaria  (Schaus)  (comb,  n.),  which  he 
collected  in  Monte  Verde,  Costa  Rica. 

Head.  Head-capsule  bears  a  pair  of  long  horn-like  processes,  flattened  and  broad  in  heliconiaria  (Fig.  100), 
not  broad  in  nigrimacula  (Fig.  101).  (Tips  of  processes  out  of  focus  in  transparency  of  cascaria.} 
Hypognathous.  Mandibles  toothed.  Six  ocelli  present.  Secondary  setae  present. 

Thorax  and  abdomen.  Not  pigmented.  Secondary  setae  on  dorsum  capitate  in  heliconiaria,  shorter  and 
trifurcate  in  nigrimacula;  row  of  knobbed  setae  run  along  midline  in  heliconiaria,  absent  from  nigrimacula. 
Scoli  absent.  Primary  setae  not  on  pinaculi.  True  legs  well  developed,  each  bearing  a  single  claw.  Prolegs 
present  on  abdominal  segments  III  to  VI,  and  on  X  (Fig.  19).  Plantae  of  ventral  prolegs  with  crochets 
biordinal  and  uniserial;  arranged  in  two  transverse  bands  in  heliconiaria  and  in  a  penellipse  in  nigrimacula. 
Abdominal  segment  X:  anal  plate  extended  into  a  pair  of  furcae;  anal  comb  present. 

Pupa  (Figs  20,  21) 

Known  for  five  species  (heliconiaria,  notes  and  illustrations  by  Kendall  (1976),  and  personal  examination; 
tipulata,  exuvial  remains;  M.  lucivittata  (Walker)  (comb,  n.),  nigrimacula,  and  cascaria  (well-preserved 
pupal  exuviae). 

The  pupa  is  rather  flattened,  although  raised  protuberances  arise  from  the  thorax  of  heliconiaria, 
lucivittata,  and  nigrimacula,  and  from  the  abdomen  of  lucivittata,  nigrimacula,  and  cascaria. 

The  pupa  of  all  five  species  is  attached  to  the  substrate  by  a  fine  silk  girdle  around  the  thorax,  and  by  a 
well-developed  cremaster.  There  is  no  cocoon.  The  temporal  cleavage  line  appears  to  be  absent,  although 
there  is  a  very  slight  indication  of  a  line  running  transversely  across  the  top  of  the  head.  In  each  case  (except 
for  heliconiaria  where  a  specimen  was  stored  in  alcohol)  the  pupa  was  attached  to  a  leaf.  However,  as  far  as 
I  know,  pupation  occurred  in  captivity  in  all  specimens  available  so  whether  or  not  hedylid  pupae  are 
always  attached  to  leaves  in  the  wild  requires  confirmation. 

Host-plant 

Known  only  for  heliconiaria:  Buettneria  aculiata  Jacq.,  (Sterculiaceae).  Recorded  by  Kendall  (1976). 

Habits 

The  following  comments,  on  heliconiaria,  are  based  on  Kendall  (1976),  who  has  provided  what  appears  to 
be  the  only  published  information.  The  eggs  are  laid  singly  on  top  of  leaves  of  Buettneria  aculiata.  The  larva 
eats  a  series  of  small  holes  in  the  leaf.  At  rest  it  lies  along  the  midrib.  Kendall  notes  that  the  adults  of 
heliconiaria  are  diurnal,  as  opposed  to  nocturnal,  which  may  be  exceptional  for  hedylids.  Similar 
observations  were  made  for  the  larva  of  cascaria  collected  by  T.  Fox  from  Costa  Rica  (pers.  comm.).  The 
specimen  was  found  lying  along  the  midrib  on  the  upper  surface  of  the  leaf.  The  leaf  of  the  food-plant 
(unidentified)  to  which  the  pupal  exuviae  is  attached  is  largely  consumed.  From  what  remains  it  is  clear  that 
the  larva  had  eaten  several  holes  in  the  leaf. 


260 


M.  J.  SCOBLE 


Parasites 

Mirax  sp.  and  Apanteles  sp.  (Braconidae),  identified  by  P.  M.  Marsh,  USDA,  Washington  D.C.,  have 
been  reared  from  heliconiaria  larvae  (R.  O.  Kendall,  pers.  comm.). 

Distribution 

Found  in  Central  and  South  America:  recorded  from  Paraguay,  Brazil,  Bolivia,  Peru,  Ecuador,  Colombia, 
Venezuela,  Guyana,  Surinam,  French  Guiana,  Panama,  Costa  Rica,  Honduras,  Guatemala,  British 
Honduras,  Mexico,  Cuba,  and  Trinidad. 

Specimens  have  been  collected  mostly  from  montane  rain  forest,  occurring  in  many  localities  in  the 
Andes.  Material  has  been  collected  at  localities  up  to  7000  ft. 

Comments  on  selected  structures 

Adult 

In  this  section  further  detail  is  provided  about  structures  selected  for  their  biological  or  systematic  interest. 
Emphasis  has  been  placed  on  comparing  the  Hedylidae  with  other  macrolepidopterans,  and  in  particular 
the  Rhopalocera.  I  include  as  macrolepidopterans  the  following  taxa:  Castnioidea,  Hesperioidea,  Papi- 
lionoidea,  Geometroidea  (sensu  Common,  1970),  Calliduloidea,  Bombycoidea  (including  Sphingidae  and 
Saturniidae),  and  Noctuoidea.  (Geometroidea  is  used  as  a  term  of  convenience  in  this  work.  Minet  (1983) 
argued  that  the  superfamily  is  not  monophyletic.) 

Head.  The  compound  eyes  are  large  and  constitute  a  relatively  large  portion  of  the  total  area  of  the  head 
(Figs  22-24).  There  is  some  variation  in  size  between  species.  The  inner,  dorsal  angle  of  each  eye  is  weakly 
emarginate  where  it  meets  the  base  of  the  antenna  (Fig.  22).  Eye-size  is  of  little  value  in  assessing 
phylogenetic  relationships  of  higher  taxa  since  it  is  related  to  habits.  Large  eyes  tend  to  be  associated  with 
low  light  intensity.  However,  the  large  eyes  of  most  true  butterflies,  a  group  that  is  typically  diurnal,  'are 
specialized  for  high  visual  acuity  at  the  expense  of  relatively  poor  absolute  sensitivity'.  The  reverse  is 
apparently  true  of  moths  (Bernard,  quoted  in  Ferguson,  1985).  Most  hedylids  are  apparently  nocturnal;  on 
independent  occasions,  specimens  have  been  collected  at  light  by  A.  Watson,  M.  J.  Matthews,  and  V.  O. 
Becker  (pers.  comm.),  and  by  the  late  C.  L.  Collenette  (label  data).  However,  heliconiaria  is  said  to  be 
diurnal  by  Kendall  (1976). 


22  23 

Figs  22,  23     Head  of  Macrosoma  tipulata  Hiibner.  22,  anterior  aspect;  23,  lateral  aspect. 


The  narrow  frons  bulges  moderately  (Fig.  23),  but  not  to  the  degree  found  for  example  in  the 
Heliothinae  (Noctuidae).  The  narrowness  of  the  frons  means  that  the  bases  of  the  antennae  are  close 
together  (Fig.  22).  The  antennae  are  usually  filiform  (Figs  26,  27),  but  in  some  species  they  are  bipectinate 
(Fig.  25).  In  those  with  filiform  antennae,  the  flagellar  segments  are  rectangular,  sometimes  almost  square, 
and  each  bears  various  sensilla.  Usually  the  sensilla  are  long  and  arranged  approximately  in  the  shape  of  a 
'U'  on  each  side  of  the  segments.  They  are  shorter  in  females.  In  tipulata  the  segments  are  uniformly 
covered  with  shorter  sensilla  (Fig.  27).  The  more  distal  flagellar  segments  bear  two  or  three  longer  setae, 
and  also  a  sensillum  basiconicum. 


NEW  CONCEPT  OF  BUTTERFLIES 


261 


25 

Figs  24-29  Head  and  associated  structures  of  Hedylidae.  24,  head,  lateral  aspect,  of  Macrosoma 
hyadnthlna  (Warren),  x  7-4;  25,  antenna  of  M.  semlermis  (Prout),  d";  26-28,  antennal  segments  of  (26) 
M.  hyacinthina(  Warren);  (27)  M.  tipulataH\ibner;(28)M.  venodes(Guen€e),  $,  arrow  indicates  reduced 
pectination;  (29)  labial  palpus  of  M.  tipulata  Hiibner. 

Bipectinate  antennae  are  found  in  both  sexes  of  Macrosoma  (=  Hedyle)  heliconiaria,  and  its  close 
relatives  M.  semiermis  (Prout)  (comb,  n.),  M.  inermis  (Prout)  (comb,  n.),  and  M.  albipannosa  (Prout) 
(comb,  n.),  and  in  males  of  Macrosoma  (=  Venodes)  napiaria  (comb.  n.).  In  the  females  of  napiaria  each 
pectination  is  reduced  to  a  small  knob  on  each  side  of  every  flagellar  segment  (Fig.  28).  In  those  species 
with  bipectinate  antennae  each  pectination  bears  sensilla.  The  sensilla  in  males  are  slightly  longer  than  in 
females. 


262  M.  J.  SCOBLE 

The  antennae  extend  about  one-half  to  just  under  two-thirds  of  the  length  of  the  fore  wing.  Whether 
filiform  or  bipectinate  they  bear  lamellar  scales  dorsally. 

Chaetosemata  are  present,  one  chaetosema  behind  each  eye  (Fig.  22).  They  are  fairly  small,  but  can  be 
seen  on  dried  specimens.  Ocelli  are  absent.  The  presence  of  chaetosemata  and  the  absence  of  ocelli  are 
conditions  found  in  Hesperioidea,  Papilionoidea,  and  in  many  Geometroidea  amongst  macrolepidopter- 
ans.  Chaetosemata  are  absent  from  the  front  of  the  head.  According  to  Jordan  (1923),  most  Hesperiidae 
have  a  pair  of  chaetosemata  additional  to  those  on  the  vertex,  sited  between  the  crest  of  scales  on  the  frons 
and  the  antennae.  This  additional  pair  is  absent  from  papilionoids. 

Mouthparts.  The  maxillary  palpi  are  minute  (Figs  22,  23),  a  condition  found  generally  in  larger 
lepidopterans.  The  proboscis  is  well  developed.  Its  base  is  not  scaled,  which  contrasts  with  the  typically 
scaled  condition  found  in  Pyralidae.  The  labial  palpus  (Fig.  29)  is  three-segmented.  The  palpi  are  generally 
moderately  ascending  (Fig.  24)  as  a  result  of  curvature  of  the  basal  segment,  but  are  not  appressed  to  the 
head  as  they  are  in  papilionids  and  pierids.  The  invaginated  sensory  pit  of  the  terminal  segment  is  a 
character  of  the  lepidopteran  ground  plan  (Kristensen,  1984ft).  In  those  hedylids  examined  for  this 
character,  the  pit  is  deep,  or  moderately  deep,  and  narrow. 

Vestiture.  Long  narrow  scales  form  the  main  covering  of  the  frons  and  vertex  (Fig.  24).  These  scales  are 
neither  closely  appressed  to  the  head,  nor  do  they  form  a  strongly  erect  tuft.  The  labial  palpi  are  covered 
with  scales;  on  the  lower  surfaces,  particularly  of  the  two  proximal  segments,  they  are  long  and  narrow, 
forming  a  fringe  (Fig.  24). 

Thorax.  To  judge  from  the  distribution  of  the  sclerotized  and  membranous  conditions  of  the  patagia  in  true 
butterflies  (Ehrlich,  1958ft),  the  membranous  condition  must  have  arisen  independently  on  several 
occasions.  The  sclerotized  state  is  regarded  as  primitive.  The  development  of  membranous  patagia  has 
probably  occurred  independently  in  other  macrolepidopterans,  although  the  general  condition  (including 
that  of  hesperiids)  is  of  sclerotized  patagia.  The  loss  of  sclerotized  patagia  is  given  as  a  butterfly  'trend'  by 
Ehrlich  (1958ft). 

The  size  of  the  anepisternum  of  the  mesothorax  (Fig.  30)  is  not  reduced,  i.e.,  there  is  no  pronounced 
dorsal  movement  of  the  anapleural  cleft.  A  reduction  is  a  prominent  and  specialized  feature  of  Papi- 
lionoidea (Brock,  1971;  figures  of  Ehrlich,  1958ft).  In  Hesperiidae  the  reduction  is  not  so  pronounced. 
Scott  (1985)  states  that  it  is  very  large  in  moths  and  skippers;  however,  it  is  not  large  in  all  skippers. 

In  the  mesothorax,  the  precoxal  sulcus  of  hedylids  appears  to  be  fused  with  the  sulcus  (Fig.  31)  called 
'marginopleural'  by  Shepard  (1930)  and  Brock  (1971).  Usually  these  sulci  are  separate  in  the  Lepidoptera. 
The  precoxal  sulcus  does  not  curve  towards  the  midline  as  it  does  in  moths  and  skippers,  and  the 
basisternum  is  'open'  -  not  'closed'  by  the  sulcus.  Brock  (1971)  drew  attention  to  the  divergent  fused 
condition  in  papilionoids  and  noted  that  the  primitive  arrangement  was  retained  in  some  Hesperiidae,  but 
almost  reached  the  specialized  condition  in  advanced  members  of  the  group.  Scott  (1985)  observed  that  in 
hesperiids,  although  the  precoxal  sulcus  (his  paracoxal  sulcus)  might  meet  the  'marginopleural',  it  does  not 
fuse  with  it  as  in  papilionoids.  These  structures  are  difficult  to  interpret  and  need  a  broader  survey  to  test 
their  soundness. 

There  are  two  characters  of  particular  importance  in  the  metathoracic  furca.  In  his  study  of  the 
integumental  anatomy  of  the  Monarch  butterfly,  Danaus  plexippus  (L.),  Ehrlich  (1958a)  noted  that  viewed 
dorsally  the  apophyses  of  the  furca  were  shaped  like  an  arrowhead.  That  this  sagittate  shape  might  be  the 
general  condition  of  both  Hesperioidea  and  Papilionoidea  was  suggested  by  Brock  (1971).  (Brock  treated 
the  Hesperiidae  as  a  family  of  the  Papilionoidea.)  In  the  Hedylidae  the  arrowhead  is  blunt,  but 
nevertheless  it  is  similar  to  that  of  other  butterflies.  The  sagittate  shape  was  not  seen  in  the  various 
macrolepidopterans  examined  by  Brock.  The  shape  is  quite  different  in  Oenochroma  vinaria  Guenee  and 
O.  polyspila  Lower  (Geometridae),  andJosiafornax  Druce  (Dioptidae),  which  were  examined  during  the 
course  of  this  work. 

The  character  of  the  metathoracic  furca  considered  by  Brock  (1971)  to  be  the  most  prominent  and 
divergent  for  the  butterflies  was  that  which  he  referred  to  as  'the  peculiar  mesal  fusion  of  the  dorsal  laminae 
of  the  secondary  arms  from  the  point  of  their  association  with  the  apophyses  to  a  point  level  with  the 
thoraco-abdominal  conjunctival  insertion'.  This  character  was  also  included  as  one  of  the  shared  derived 
(apomorphic)  features  of  the  Hesperioidea  and  Papilionoidea  by  Kristensen  (1976).  If  I  understand 
Brock's  description  correctly,  this  mesal  fusion  is  present  in  the  Hedylidae.  It  can  best  be  seen  in  dorsal 
view.  Despite  the  weight  given  to  this  character  by  Brock  (1971),  the  structure  seems  to  be  present  in 
Oenochroma  vinaria  (Geometridae:  Oenochrominae)  and  also  in  Archiearis  Hiibner,  a  member  of  the 
primitive  geometrid  subfamily  Archiearinae. 

Wing  venation  (Fig.  32a).  The  venation  does  not  seem  to  vary  between  species  either  in  the  number  of 
veins  present,  or  their  branching  pattern  and  disposition.  In  the  fore  wing  the  arrangement  of  the  veins  of 
the  radial  sector  (Rs)  is  characteristic  of  the  family.  The  sinuous  course  run  by  Rsl  and  Rs2  is  such  that  they 


NEW  CONCEPT  OF  BUTTERFLIES 


263 


a 


Figs  30-32  Hedylidae.  30,  31,  mesothorax  of  Macrosoma  rubedinaria  (Walker):  (30)  ventrolateral 
aspect,  (31)  lateral  aspect;  32,  M.  nigrimacula  (Warren)  (a)  fore  wing,  (b)  hind  wing;  anepisternum 
(am),  katepisternum  (km),  basisternum  (bm),  precoxal  sulcus  (pcs),  'marginopleural'  sulcus  (mps). 

nearly  meet  at  one  point.  The  only  stalking  that  occurs  is  that  formed  by  the  common  stem  of  Rs3  and  Rs4. 
Veins  Rl,  Rsi,  and  Rs2  are  all  separate  (at  least  from  the  cell).  These  veins  fail  to  coalesce  at  any  point  so, 
unlike  many  macrolepidopterans,  no  secondary  cells  are  formed.  This  pattern  is  generalized,  and 
approximates  to  that  exhibited  by  the  more  primitive  of  the  macrolepidopteran  elements.  Unlike  several 
lineages  of  higher  Lepidoptera,  there  is  no  inward  migration  ('splitting  back')  of  veins.  The  arrangement  of 


264 


M.  J.  SCOBLE 


33 


Figs  33-36  Wing-coupling  apparatus  in  Hedylidae.  33,  frenulum  of  Macrosoma  hyacinthina  (Warren); 
34,  retinaculum  of  M.  bahiata  (Felder),  view  from  ventral  surface;  35,  36,  frenular  bristles  of  (35)  M. 
napiaria  (Guenee),  cf,  (36)  M.  satellitiata  satellitiata  (Guenee),  $. 


Rs  therefore  provides  few  clues  to  relationships,  although  it  further  supports  the  exclusion  of  the  group 
from  advanced  geometroids,  calliduloids,  bombycoids,  and  noctuoids. 

Vein  MI  is  remote  from  Rs4,  a  condition  widely  encountered,  unlike  the  arrangement  found  in 
Epiplemidae  and  Uraniidae  where  these  branches  are  associated.  The  usual  geometroid  condition,  and 
indeed  that  of  most  higher  Ditrysia,  is  that  /?s3  and  Rs2  share  a  common  stem,  and  Rs4  arises  from  this  joint 
stem  nearer  to  the  cell  (i.e.  proximally).  A  notable  exception  is  the  Papilionoidea  where  Rsj  and  Rs4  are 
often  associated  as  in  hedylids.  In  the  Hesperioidea  there  is  no  'stalking'  of  the  veins  whatsoever;  all  run 
independently  from  the  cell  to  the  apex  of  the  termen.  Vein  M  is  very  weakly,  but  definitely,  visible  in  the 
cell. 

CuP  is  often  present,  but  weak. 

Hind  wing  venation  (Fig.  32b).  Vein  Sc+/?j  is  remote  from  Rs  from  the  base  of  the  cell;  the  condition 
contrasts  with  that  found  in  geometrids  and  drepanids,  but  is  not  dissimilar  from  Epiplemidae  and 
Uraniidae.  The  presence  of  two  anal  veins  in  the  hind  wing  is  more  frequently  encountered  in  the 
butterflies  (including  Hesperiidae),  bombycoids  and  noctuoids  amongst  the  higher  Ditrysia,  than  in 
geometroids  (s.l.).  The  rather  generalized  occurrence  of  both  these  attributes  therefore  is  of  little  value  in 
assessing  the  relationships  of  Macrosoma.  Nevertheless  both  features  indicate  yet  further  the  remoteness 
of  the  genus  from  the  Geometridae. 

M2  does  not  arise  nearer  MI  than  M3. 

Wing  coupling  (Figs  33-37).  In  males,  a  single-spined  frenulum  is  usually  well  developed  on  the  hind 


NEW  CONCEPT  OF  BUTTERFLIES 


265 


37 


Fig.  37    Macrosoma  hyacinthina  (Warren),  base  of  fore  wing,  ventral  aspect;  anterior  chamber  (ac), 
posterior  chamber  (pc),  subcostal  fold  (fd). 


wing  (Fig.  33).  It  engages  a  long  retinaculum  on  the  fore  wing  (Figs  34,  37,  39).  The  length  of  the 
retinaculum  is  related  to  the  need  for  it  to  extend  over  the  prominent  subcostal  fold  so  as  to  engage  the 
frenulum  (Figs  37, 39).  The  frenulo-retinacular  system  is  reduced  and  probably  non-functional  in  the  males 
of  three  species:  M.  subornata  (Warren)  (comb,  n.),  M.  leucoplethes  (Prout)  (comb,  n.),  and  napiaria.  In 
the  first  two  species  the  frenulum  remains  as  a  single  spine,  but  it  is  much  weaker  and  shorter.  In  napiaria  it 
is  reduced  to  a  few  short  bristles  (Fig.  35).  The  retinaculum  of  napiaria  is  lost,  while  in  subornata  and 
leucoplethes  it  is  reduced  to  a  very  short,  and  presumably  functionless  structure.  Although  the  frenulum 
and  retinaculum  are  reduced  in  subornata  and  leucoplethes,  the  system  is  well  developed  in  the  closely 
related  M.  desueta  (Prout)  (comb.  n.). 

In  females  there  are  sometimes  a  few  weak  costal  bristles  on  the  hindwing  (Fig.  36).  There  is  no  sign  of  a 
retinaculum,  and  functionally  the  frenulo-retinacular  system  is  lost  in  this  sex. 

In  Hesperioidea  and  Papilionoidea  the  frenulum  and  retinaculum  are  lost  but  remarkably,  these 
structures  are  present  in  males  of  the  hesperiid  Euschemon.  Scott  (1985)  notes  that  Euschemon  is 
otherwise  a  typical  pyrgine  hesperiid  and  suggests  an  independent  development  of  the  frenulum  and 
retinaculum  through  perhaps  a  reverse  mutation.  The  loss  of  the  frenulum  and  retinaculum  or  even  of  the 
retinaculum  alone  has  occurred  many  times  within  the  Lepidoptera  besides  the  butterflies,  e.g.  in 
bombycoids,  uraniids  and  some  drepanids. 

Wing  surface.  There  is  no  holding  area  (Haftfeld)  on  the  underside  of  the  base  of  the  fore  wing  and  the 
metascutum  of  the  thorax.  These  patches  of  velcro-like  microtrichia  (aculei)  are  absent  from  Hesper- 
ioidea, Papilionoidea,  Geometroidea,  and  most  Bombycoidea,  a  feature  considered  to  be  related  to  the 
resting  position  of  the  wings  (Common,  1969;  Kuijten,  1974).  In  general,  those  Lepidoptera  with  wings 
folded  over  the  body  have  holding  areas  on  the  fore  wings  and  thorax,  whereas  those  with  wings  held 
vertically  at  rest  or  at  right  angles  to  the  body  do  not. 

Vestiture.  Two  kinds  of  scales  predominate  on  the  wing-surface  (Fig.  38):  (a)  broad,  'typical', 
overlapping  scales,  and  (b)  hair-like  scales.  The  wings  appear  to  be  hairy  in  parts.  The  broad  scales  do  not 
provide  a  dense  cover  to  the  wings;  hedylids  are  relatively  weakly  scaled.  In  several  species  there  are  small 
hyaline  patches  devoid  of  scales,  or  larger  areas  with  semi-transparent  scales.  The  small  hyaline  patches 
may  be  seen  with  the  naked  eye  on  some  specimens  (Figs  10, 16).  The  hair-like  scales  are  more  dense  on  the 
basal  section  of  the  upperside,  and  somewhat  more  dense  on  the  underside. 

Figs  1-18  illustrates  the  main  kinds  of  wing  patterns  in  the  Hedylidae.  Of  particular  note  is  that  the 
markings  on  both  dorsal  and  ventral  surfaces  are,  with  very  minor  exceptions,  the  same,  compare  Fig.  12 
with  Fig.  18.  At  least  one  hedylid  may  hold  its  wings  vertically  at  rest  so  exposing  the  underside  (R.  O. 
Kendall,  pers.  comm.)  although  this  requires  confirmation.  Such  a  resting  position  may  be  related  to  the 
strong  patterning  on  the  underside .  In  M.  subornata  the  posterior  half  of  the  underside  is  pale  and  contrasts 


266  M.  J.  SCOBLE 

sharply  with  both  the  rest  of  the  underside  and  with  that  of  the  upper  surface.  The  habit  of  holding  the 
wings  vertically  is  common  among  butterflies,  and  is  also  to  be  seen  in  several  geometrids.  (I  am  grateful  to 
Dr  K.  Sattler  for  suggesting  the  resting  position  of  hedylids.) 

There  are  several  relatively  distinct  wing-patterns  in  hedylids,  as  well  as  other  patterns  that  are  less 
distinct.  Frequently,  elements  of  one  subgroup  are  found  in  another.  For  example,  large  areas  covered  by 
semi-transparent  scales  are  present  in  many  species  as  are  small,  but  distinctive,  white  triangular  marks  on 
the  costa  of  the  fore  wing  near  the  apex.  In  some  species  the  apex  of  the  fore  wing  bears  a  large  patch  of 
either  chestnut  or  dark  brown.  The  illustrations  show  the  distinctive  emargination  of  the  apex  of  the 
forewing,  a  condition  found  in  many  species,  and  the  weakly  falcate  shape  of  them  all.  The  comments  that 
follow  are  an  adjunct  to  the  black  and  white  photographs  to  provide  a  guide  to  colour. 

M.  heliconiaria  (Guenee)  (Fig.  1).  Ground  colour  darkish  grey-brown  with  translucent  patches  on  fore 
and  hind  wings.  The  small  triangular  mark  on  the  costa  of  the  fore  wing  is  white. 

M.  hyacinthina  (Warren)  (comb,  n.)  (Fig.  2).  Ground  colour  dark  grey-brown  with  a  purple  iridescence. 
More  brown  than  grey  at  apex  of  fore  wing.  The  pale  patches  on  both  wings  are  covered  with 
semi-transparent  scales  except  for  the  subtriangular  mark  on  the  costa  of  the  fore  wing,  which  is  white. 

M.  tipulata  Hiibner  (Fig.  3).  A  pale  species.  The  wings  bear  large  areas  of  semi-transparent  scales.  The 
darker  areas  are  pale  brown,  and  the  small  irregular  patches  on  the  fore  wing  are  white,  edged  with  darker 
brown. 

M.  rubedinaria  (Walker)  (comb,  n.)  (Fig.  4).  Ground  colour  brown.  The  small  dark  streaks  are  dark 
brown,  edged  with  white. 

M.  ustrinaria  (Herrich-Schaffer)  (comb,  n.)  (Fig.  5).  A  pale  species.  The  darker  scales  scattered 
unevenly  over  the  wing  surfaces  are  pale  brown. 

M.  leucophasiata  (Thierry-Mieg)  (comb,  n.)  (Fig.  6).  The  pale  areas  are  white  and  the  dark  parts  dark  to 
very  dark  grey-brown. 

M.  subornata  (Warren)  (Fig.  7).  Ground  colour  brown,  dark  brown  near  apices  of  wings.  The  patch  on 
the  costa  is  cream,  and  the  patch  below  it  and  the  tiny  spot  to  one  side  are  white.  On  the  underside  the 
posterior  half  of  the  fore  wing  is  pale,  contrasting  markedly  with  the  equivalent  section  of  the  upper 
surface. 

M.  paularia  (Schaus)  (comb,  n.)  (Fig.  8).  Ground  colour  of  fore  wing  pale  grey-brown  flecked  and 
streaked  with  dark  brown.  The  large  spot  at  the  apex  of  the  fore  wing  is  a  warm  pale  brown.  The  large  pale 
area  on  the  hind  wing  is  composed  of  semi-transparent  scales. 

M.  coscoja  (Dognin)  (comb,  n.)  (Fig.  9).  Ground  colour  pale  brown.  Apex  of  fore  wing  dark  brown.  The 
small  patch  at  the  base  of  the  strong  apical  patch  is  composed  of  semi-transparent  scales. 

M.  satellitiata  (Guenee)  (comb,  n.)  (Fig.  10).  Ground  colour  a  darker  brown  than  that  of  coscoja. 
Otherwise  similar  to  that  species.  The  small  'white'  spot  at  the  base  of  the  hind  wing  represents  the  glassy 
modification  in  males  discussed  below. 

M.  albifascia  albifascia  (Warren)  (comb,  n.)  (Fig.  11).  Ground  colour  grey-brown.  The  pale  area  on  the 
fore  wing  is  semi-transparent.  The  apex  of  the  fore  wing  is  a  warm  pale  brown  thickly  edged  with  dark 
brown.  The  very  dark  streak  and  patches  on  the  fore  wing  are  also  dark  brown. 

M.  hedylaria  (Warren)  (comb,  n.)  (Figs  12, 18).  The  colour  pattern  is  extremely  similar  to  heliconiaria, 
although  the  apex  of  the  fore  wing  is  more  brown  than  grey-brown.  The  name  hedylaria  is  obviously  a 
composite  of  Hedyle  (the  genus  in  which  heliconiaria  was  described  but  which  is  treated  as  a  junior 
synonym  of  Macrosoma  in  the  present  work)  and  heliconiaria. 

M.  lucivittata  (Walker)  c?  (Fig.  13).  Ground  colour  brown.  The  pale  areas  are  covered  with  semi- 
transparent  scales  and  many  brown  piliform  scales.  9  (Fig.  14).  Similar  to  the  male  but  with  a  distinct 
semi-transparent  spot  near  the  apex  of  the  fore  wing. 

M.  nigrimacula  (Warren)  (Fig.  15).  Ground  colour  brown,  flecked  and  spotted  with  dark  brown.  The 
semi-transparent  areas  are  poorly  defined  on  the  upper  surface  in  this  species.  On  the  underside  of  the  fore 
wing  the  posterior  half  bears  a  large  pale  grey  area  that  contrasts  with  the  rest  of  the  wing. 

M.  leptosiata  (Felder)  (comb,  n.)  (Fig.  16).  Ground  colour  brown.  Both  fore  and  hind  wing  edged  with 
darker  brown  distally,  particularly  on  the  fore  wing. 

M.  napiaria  (Guenee)  (Fig.  17).  Ground  colour  off-white,  formed  by  a  covering  of  semi-transparent 
scales.  The  veins  of  the  fore  wing  are  a  more  strongly  pigmented  shade  of  cream. 

Modification  of  the  fore  wings  (Figs  37, 39-43).  The  base  of  the  fore  wing  bears  some  apparently  unique 
modifications.  These  take  the  form  of  a  ventral  expansion  of  vein  Sc  into  a  fold,  and  the  presence  of  two 
small,  rounded  chambers  developed  within  the  bases  of  certain  veins.  A  pocket  is  also  present  and  is 
enclosed  by  a  dense  fringe  of  scales.  These  modifications  occur  within  both  males  and  females.  In  addition, 
a  distinct  lobe  is  formed  near  the  anal  edge  of  the  fore  wing  in  males  of  some  species. 


NEW  CONCEPT  OF  BUTTERFLIES 


267 


Figs  38-43  Fore  wing  structure  (ventral  aspects)  of  Hedylidae.  38,  vestiture  of  fore  wing  of  Macrosoma 
tipulata  Hiibner;  39,  base  of  fore  wing  of  M.  hyacinthina  (Warren)  to  show  subcostal  fold  (fd);  40,  41, 
scanning  electron  micrographs  of  base  of  fore  wing  of  M.  tipulata  Hiibner;  (40)  x  16,  large  arrow 
indicates  pocket,  small  arrow  shows  base  of  retinaculum,  (41)  detail  of  pocket  x  40,?  tympanum  (tm); 
42,  base  of  fore  wing  of  M.  hyacinthina  (Warren),  arrow  indicates  position  of  membrane  (see  text);  43, 
lobe  at  base  of  fore  wing  of  M.  hyacinthina  (Warren). 


268  M.  J.  SCOBLE 

The  expansion  of  Sc  on  the  underside  of  the  fore  wing  is  prominent  (Figs  37,  39,  40).  There  is  some 
variation  in  its  extent  between  species,  but  the  form  is  basically  similar.  This  fold  is  a  hollow  outgrowth  of 
the  vein  (although  both  membranes  are  close  together  and  not  'ballooned').  The  fold  extends  a  short  way 
along  the  wing.  At  the  extreme  base  of  the  fore  wing,  Sc  is  expanded  into  a  hollow  chamber  (Figs  37, 42)  the 
main  part  of  which  lies  ventral  to  the  plane  of  the  wing.  This  chamber  is  divided  from  the  fold  by  a 
membrane  that  runs  at  right  angles  to  the  plane  of  the  wing.  The  position  of  this  membrane  is  arrowed  in 
Fig.  42;  it  is  visible  as  a  dark  line  in  this  view.  The  chamber  is  invaginated  caudally  effectively  providing 
another  pocket,  which  is  enclosed  by  a  thick  fringe  of  hair-scales  (Figs  40,  41).  The  posterior  wall  of  the 
chamber  is  invaginated.  From  the  ventral  exterior  lip  of  this  invagination  arises  a  thick  fringe  of  hair-scales 
(Fig.  40  and  particularly  Fig.  41).  Below  this  fringe,  and  presumably  protected  by  it,  is  an  extremely 
delicate,  transparent  membrane.  This  is  supported  by  a  cuticular  frame  at  the  base  of  Cu,  also  lying  in  the 
ventral  plane  of  the  wing.  Stretched  over  the  other  (posterior)  side  of  this  frame  is  a  whitish  membrane, 
which  closely  resembles  the  tympanum  found  in  the  auditory  organs  of  certain  Lepidoptera  (Fig.  41). 

Lying  mainly  within  the  dorsal  plane  of  the  fore  wing  is  another  small ,  closed  chamber  (Figs  37 , 42) .  This 
chamber  is  an  expansion  of  the  base  of  the  cubitus,  'Cu^  in  the  terminology  of  Sharplin  (1963). 


47 

.«->ja  ?? 


Figs  44-47  Scanning  electron  micrographs  of  specialized  structures  on  hind  wing  (ventral  aspects)  of 
Hedylidae.  44-46,  Macrosoma  tipulata  Hiibner:  (44)  general  view  of  area  x  16,  (45)  detail  of  squat 
protuberance,  x  140,  (46)  detail  of  finger-like  protuberance,  x  160;  47,  M.  zikani  conifera  (Warren) 
(comb,  n.),  base  of  hind  wing  x  16. 


NEW  CONCEPT  OF  BUTTERFLIES 


269 


50 


51 


Figs  48-51  Wing  structures  of  Hedylidae.  48,  49,  scale  from  specialized  area  of  hind  wing  (ventral 
surface)  ofMacrosoma  tipulata  Hiibner,  (48)  whole  scale,  x  2600,  (49)  detail  to  show  serrations  x  6500; 
50,  specialized  structure  near  base  of  hind  wing  (ventral  surface)  of  M.  nigrimacula  (Warren);  51,  base  of 
fore  wing  of  M.  hyacinthina  (Warren)  to  show  position  of  second  median  plate  partly  hidden  under  \A 
(arrowed). 

Although  the  function  of  these  chambers  is  unknown,  there  are  several  similarities  with  the  tympanal 
organs  found  at  the  base  of  the  fore  wing  in  many  nymphalid  butterflies.  The  structure  of  these  alar 
tympanal  organs  is  summarized  by  Bourgogne  (1951).  In  both  hedylids  and  those  nymphalids  with  these 
organs,  the  tympanum  is  situated  at  the  base  of  vein  Cu.  Unlike  thoracic  or  abdominal  hearing  organs, 
there  is  no  tympanic  cavity;  instead,  the  tympanum  is  superficial.  In  nymphalids,  the  membrane  is 
protected  by  scales,  but  in  hedylids  it  is  more  exposed.  In  most  of  those  nymphalids  with  alar  hearing 
organs,  veins  Sc  and  Cu  are  swollen  basally,  and  the  tracheal  air  sacs  within  the  swellings  communicate  with 
the  tympanic  sac  (Bourgogne,  1951).  It  is  uncertain  whether  there  is  a  functional  relationship  between  the 
sacs,  but  it  should  be  noted  that  not  all  nymphalids  with  hearing  organs  have  swollen  veins.  In  the 
Hedylidae  the  veins  are  not  swollen. 

It  is  well  known  that  some  butterflies  respond  to  sound.  Swihart  (1967)  demonstrated  the  sensitivity  to 


270 


M.  J.  SCOBLE 


sound  of  a  small  chamber  at  the  base  of  each  hind  wing  of  Heliconius  erato  (L.).  He  also  found  a  similar,  but 
smaller,  chamber  at  the  base  of  each  fore  wing. 

No  information  is  available  on  the  question  of  whether  hedylids  respond  to  sound,  but  on  circumstantial 
evidence  it  seems  likely  that  the  modifications  at  the  base  of  the  fore  wing  function  as  an  auditory  organ. 

The  possibility  that  the  chamber  enclosed  by  the  fringe  of  hairs  is  a  scent  pouch  cannot  be  discounted ,  but 
the  pocket  is  present  in  both  males  and  females.  The  presence  of  'scent  pockets'  tends  to  be  sexually 
dimorphic  in  lepidopterans,  but  is  not  always  so. 

A  small  lobe  extends  from  the  anal  edge  of  the  fore  wing  near  its  base  in  the  males  of  some  species.  It  is 
prominent  in  hyacinthina  (Fig.  43).  A  very  weakly  developed  expansion  of  this  area  is  found  in  many 
species. 

Modifications  of  the  hind  wing  (males).  These  are  present  in  many  species.  Viewed  with  the  naked  eye 
they  appear  as  small,  oval,  glassy  areas  at  the  base  of  the  wing  on  both  the  upper  and  the  under  sides  (Figs 
10,  16).  The  modifications  take  the  form  of  a  folding  of  the  wing  membrane,  making  the  surface  appear 
rather  crinkled  or  distorted  when  viewed  under  a  dissecting  microscope.  Their  appearance  under  the 
scanning  electron  microscope  is  shown  in  Figs  44  and  47.  From  this  area  one  or  more  protuberances  arise  on 
the  under  side  of  the  wing  (Figs  44-47,  50).  The  area  is  covered  with  specialized  scales. 

In  hyacinthina  the  modified  area  may  be  roughly  divided  into  an  inner,  approximately  oval  section, 
which  bears  short  piliform  scales  on  both  surfaces,  and  an  outer  section  with  both  piliform  scales  and 
semi-transparent  laminar  scales.  A  prominent  knob-like  structure  is  present  on  the  underside. 

The  modifications  of  the  hind  wing  are  well  developed  in  many  other  species,  and  less  pronounced  in 
several  more.  Modifications  present  in  one  species  may  be  absent  from  closely  related  ones.  In  M.  bahiata 
(Felder)  (comb,  n.)  for  example  they  are  present,  while  in  coscoja,  a  close  relative,  they  are  not.  In  the 
same  group  of  species  the  structures  are  less  strongly  pronounced  in  M.  albistria  (Prout)  (comb,  n.)  and  M. 
uniformis  (Warren).  In  tipulata  two  protuberances  are  present  (Fig.  44)  of  which  one  is  relatively  squat 
(Fig.  45),  and  the  other  is  a  long  digitate  structure,  which  is  slightly  swollen  apically  (Fig.  46).  The  function 
of  these  protuberances  is  uncertain,  but  since  they  are  confined  to  the  males  they  are  probably  scent  organs 
of  some  kind.  Figs  48  and  49  illustrate  the  specialized  scales  found  on  the  surface  of  the  structure  in  tipulata. 
The  longitudinal  ridges  are  strongly  serrated  (Fig.  49). 


54 


Figs  52-54    Fore  legs  of  male  Hedylidae.  52,  Macrosoma  semiermis  (Prout),  tibia  and  tarsus;  53,  54, 
epiphysis  of  (53)  M.  tipulata  Hiibner,  (54)  M.  napiaria  (Guenee). 


NEW  CONCEPT  OF  BUTTERFLIES  271 

Wing  base.  According  to  Dr  R.  de  Jong  (pers.  comm.  and  in  prep.),  to  whom  I  am  indebted  for  sketches 
and  information,  the  second  median  plate  of  the  fore  wing  is  always  partly  hidden  under  the  base  of  vein  1A 
in  both  Hesperioidea  and  Papilionoidea.  The  general  ditrysian  condition  seems  to  be  that  the  plate  is  not 
obscured  by  1A.  In  the  Hedylidae  the  second  median  plate  is  small  and  partly  obscured  by  1A  (Fig.  51). 

Legs.  An  epiphysis  is  present  on  the  fore  leg  of  both  males  and  females  (Figs  52-56).  The  presence  of  this 
structure  is  widespread  among  macrolepidopterans  including  the  Hesperiidae  (with  a  few  exceptions)  and 
Papilionidae,  but  it  is  lost  in  many  Pieridae,  Nymphalidae,  Libytheidae,  and  Lycaenidae. 

In  males  the  fore  tarsi  are  modified  (Figs  55, 58).  Although  they  are  not  reduced  in  length  (in  fact  they  are 
long  or  very  long)  the  segments  are  mostly  fused  so  that  only  a  long  proximal  segment  and  a  short  distal 
segment  are  present.  In  tipulata  the  proximal  segment  is  extremely  long  (Fig.  55). 

The  pretarsus  of  the  fore  leg  is  reduced  to  a  pair  of  small  claws  in  males  (Figs  52,  58).  In  some  species 
these  claws  are  minute,  and  in  all  species  they  are  only  very  weakly  curved.  There  is  no  arolium,  or 
empodium,  nor  are  there  pulvilli  on  the  fore  tarsi  of  males.  The  fore  tarsi  bear  a  prominent  fringe  of 
piliform  scales  giving  them  a  feathery  appearance.  There  are  no  spines. 

In  females  the  fore  tarsi  are  not  modified  (e.g.  tipulata,  Fig.  56).  There  are  five  tarsomeres  and  the 
pretarsus  bears  claws  that  are  strongly  curved  and  asymmetrically  forked,  pulvilli,  and  a  large  arolium.  The 
tarsomeres  are  covered  with  laminar,  not  piliform,  scales,  and  a  series  of  small  spines  is  present. 

The  mid  and  hind  tarsi  in  both  males  and  females  are  five-segmented,  spined,  and  with  a  normally 
developed  pretarsus  with  claws,  pulvilli  and  an  arolium.  The  mid  leg  is  generally  longer  than  the  hind  leg, 
but  shorter  than  the  fore  leg. 

Microtrichia  or  aculei  ('tiny  spines'  of  Scott,  1985)  are  not  present  on  the  dorsal  aspect  of  the  tarsi,  unlike 
the  Papilionidae,  Pieridae,  some  Nymphalidae  and  some  Lycaenidae.  According  to  Scott  they  are  absent 
from  hesperiids  and  from  other  macrolepidopterans.  Microtrichia  are,  however,  present  on  the  ventral 
aspect  of  the  tarsi  of  Hedylidae  although  their  distribution  is  patchy,  being  best  developed  on  the  pulvilli. 
They  are  absent  from  the  fore  tarsi  of  males.  Ventral  microtrichia  are  probably  widespread  in  Lepidoptera. 

A  single  pair  of  tibial  spurs  is  present  on  the  midleg  of  both  sexes  (Fig.  61).  The  hind  leg  of  the  male  of 
tipulata  bears  two  pairs  of  tibial  spurs  (Fig.  62),  but  usually  hedylids  (including  the  female  of  tipulata)  have 
one  pair  distally  on  the  hind  tibia.  The  proximal  pair  of  spurs  is  found  in  many  Lepidoptera,  e.g.  some 
Thyrididae,  some  Hesperiidae,  some  Drepanidae,  many  Bombycoidea  (including  Sphingidae),  and  many 
Noctuoidea  (including  Notodontidae).  This  loss  is  a  classic  example  of  a  character  that  is  prone  to 
independent  reduction  in  many  groups.  Therefore  the  loss  of  spurs  in  most  hedylids  and  in  papilionoids 
does  not  provide  strong  evidence  for  their  monophyly.  In  one  female  specimen  of  hyacinthina  there  is 
apparently  no  pretarsus  at  all  on  the  hind  leg:  the  tarsus  ends  in  a  flask-shaped  tarsomere  (Fig.  60).  There  is 
no  sign  of  claws  arising  from  this  last  tarsomere.  Of  the  two  other  females  in  the  collection  of  the  British 
Museum  (Natural  History),  one  has  lost  both  hind  legs  while  in  the  other  the  tarsi,  including  the  pretarsus, 
are  normal.  In  many  species  the  hind  tibia  is  slightly  swollen. 

A  scent  pencil  occurs  on  the  fore  tibia  of  males  of  five  species  (hyacinthina,  lucivittata,  M.  klagesi  (Prout) 
(comb,  n.),  and  nigrimacula,  and  probably  (only  one  specimen  available)  on  M.  latiplex  (Dognin)  (comb. 
n.)).  The  pencil  comprises  a  bunch  of  long  hairs  arising  from  the  inner  surface  of  the  tibia.  It  is  particularly 
well  developed  in  hedylaria. 

The  modifications  of  the  legs  of  the  Hedylidae  are  all  found  within  other  butterflies.  Information  for 
Hesperioidea  and  Papilionoidea  is  summarized  in  a  useful  table  by  Scott  (1985:  table  1).  Pulvilli  and  the 
arolium  are  lost  from  the  legs  of  Papilionidae,  some  Pieridae,  and  some  Nymphalidae.  In  the  Papilionidae 
and  the  Pieridae  the  tarsomeres  are  not  fused.  In  Nymphalidae  the  male  fore  legs  are  small  although  the 
tarsomeres  are  not  lost  or  fused.  In  Lycaenidae  the  male  fore  legs  are  reduced  to  a  single  tarsomere 
together  with  reduction  or  loss  of  claws.  The  proximal  pair  of  tibial  spurs  on  the  hind  leg  is  absent  from  all 
true  butterflies  (Ehrlich,  1958ft;  Scott,  1985)  and  hedylids  (with  the  exception  of  the  male  of  tipulata),  but 
these  spurs  are  present  in  skippers.  Scent  brushes,  when  present,  tend  to  be  found  on  the  hind  legs  of 
butterflies;  they  occur  on  the  fore  legs  of  Hedylidae. 

Abdomen.  The  abdomen  is  distinctly  curved,  particularly  in  males  (Fig.  63),  and  is  laterally  flattened.  Not 
only  are  these  features  butterfly-like,  but  from  my  initial  observations  they  resemble  true  butterflies 
(Papilionoidea)  rather  than  skippers  (Hesperioidea).  This  interpretation,  however,  remains  tentative. 

Tergum  I  is  'pouched'  (Figs  64,  68),  that  is,  there  is  a  pocket-like  outgrowth  of  tergum  I  posteriorly  that 
overlaps  tergum  II.  Ehrlich  (19586)  used  the  word  'pouched'  to  describe  the  typical  papilionoid  condition. 
The  pouching  in  hesperioids  is  not  strong,  and  in  some  papilionids  the  pouch  is  reduced  (Ehrlich,  1958ft). 

On  segment  I  there  are  pre-  and  postspiracular  bars.  The  prespiracular  bar  (Figs  64,  68)  is  a  narrow 
sternopleural  structure,  which  extends  anteriorly  from  the  antero-lateral  margin  of  sternum  II,  and  curves 
around  the  spiracle.  The  presence  of  a  prespiracular  bar  is  the  general  condition  in  macrolepidopterans, 
including  hesperioids  and  papilionoids,  although  it  is  lost  in  pierids  (Ehrlich,  1958ft).  The  presence  of  this 


272 


M.  J.  SCOBLE 


N 


•*. 


55 


56 


57 


58 


Figs  55-58     Fore  legs  of  Hedylidae.  55,  56,  Macrosoma  tipulata  Hiibner:  (55)  cf ,  (56)  $ ;  57,  M.  hedylaria 
(Warren),  O";  58,  M.  semiermis  (Prout),  tarsal  claws  of  O",  arrowed. 


NEW  CONCEPT  OF  BUTTERFLIES 


273 


59 


60 


61 


Figs  59-62  Legs  of  Hedylidae.  59,  pretarsus  of  Macrosoma  tipulata  Hiibner,  9,  showing  weak  fork  of 
claw;  60,  last  (5th)  tarsal  segment  of  hind  leg  of  M.  hyadnthina  (Warren),  $ ;  61,  62,  hind  tibia  of  (61)  M. 
hedylaria  (Warren),  c? ,  (62)  M.  tipulata  Hiibner,  cf . 

structure  in  the  Hedylidae  is  therefore  of  no  particular  phylogenetic  significance.  Brock  (1971)  considers 
that  both  pre-  and  postspiracular  bars  are  secondarily  developed  within  the  Papilionoidea  and  Hesper- 
ioidea  (his  Papilionoidea)  and  analogous  to  similar  structures  in  certain  Pyraloidea,  Geometroidea  and 
Noctuoidea.  It  is  not  entirely  clear  why  Brock  does  not  consider  them  homologous. 

The  postspiracular  bar  is  well-developed  in  the  Hedylidae.  It  is  a  tergopleural  structure  (Figs  64,  68), 
which  just  fails  to  meet  the  sternum.  According  to  Brock  (1971)  the  postspiracular  bar  is  derived  from  terga 
I  and  II.  Ehrlich  (19586)  considered  the  presence  of  a  postspiracular  bar  to  be  specialized  within  the 
Papilionoidea,  where  he  presumably  considers  it  to  have  evolved  independently  within  the  group.  Scott 
(1985)  records  the  bar  as  absent  in  Papilionidae  (Ehrlich  said  it  was  reduced  or  absent),  present  or  absent  in 
Nymphalidae,  present  and  large  in  Libytheidae,  and  absent  in  Lycaenidae.  He  notes  its  presence  in  or 
absence  from  Hesperioidea  and  its  absence  from  other  macrolepidopterans.  Both  Ehrlich  and  Scott 
consider  the  structure  as  derived.  The  occurrence  of  a  similar  structure  in  some  other  macrolepidopterans 


274 


M.  J.  SCOBLE 


63 


64 


66 


tg  Vlll 


67 


65 


Figs  63-67  Abdomen  of  Hedylidae.  63,  64,  Macrosoma  bahiata  (Felder),  lateral  aspect:  (63)  whole 
abdomen,  (64)  base  of  abdomen,  small  arrow  -  pre  spiracular  sclerotization,  large  arrow  -  post 
spiracular  sclerotization;  65,  sternum  II  (st  II)  of  M.  paularia  (Schaus);  66,  segment  VIII  of  M.  semiermis 
(Prout),  d",  lateral  aspect;  67,  tergum  VIII  (tg  VIII)  of  M.  tipulata  Hubner,  cf . 


(e.g.  Thyrididae)  noted  by  Brock  is  probably  too  isolated  to  suggest  that  the  structure  in  the  butterflies  is 
not  derived. 

The  tergal  groove  is  well  developed  and  leads  into  a  strong  tergal  brace  -  the  internal  ridge  of  the  groove 
(Fig.  64). 

There  are  no  abdominal  tympanal  organs  in  the  Hedylidae. 

Male  genitalia  (Figs  69-89).  The  lateral  arms  of  the  vinculum  are  long,  so  the  genital  capsule  appears 
'deep',  that  is,  expanded  dorso-ventrally.  The  natural  way  to  mount  these  structures  for  microscopic  slide 
preparation  is  laterally,  since  if  they  are  mounted  ventrally  they  tend  to  topple  over  on  the  slide.  Most 


NEW  CONCEPT  OF  BUTTERFLIES 


275 


68 


69 


Figs  68-71  Abdomen  and  male  genitalia  of  Hedylidae.  68,  segments  I  and  II  of  Macrosoma  bahiata 
(Felder),  lateral  aspect,  abbreviations  as  for  Fig.  64;  69-71,  male  genitalia,  lateral  view,  of  (69)  M. 
semiermis  (Prout),  (70)  M.  hyacmthina  (Warren),  (71)  M.  paularia  (Schaus). 

butterfly  systematists  mount  male  genitalia  laterally  for  the  same  reason.  The  genitalia  of  the  Hedylidae  are 
similar  to  those  of  many  butterflies  in  the  shape  of  the  genital  capsule.  They  might  be  regarded  as  fairly 
generalized  butterfly  genitalia,  but  this  does  not  provide  strong  evidence  for  the  monophyly  of  hedylids  and 
other  butterflies  since  the  arrangement  might  be  primitive.  Certainly  the  Castniidae  also  have  genitalia  that 
are  'deep'.  The  lateral  arms  are  fused  with  the  tegumen  so  that  segment  IX  of  the  abdomen  forms  a  closed 
ring  (e.g.  Figs  73,  74),  a  feature  noted  as  a  tentative  ground  plan  character  of  the  Amphiesmenoptera 
(Kristensen  &  Nielsen,  1979:  126;  Kristensen,  1984&:  150).  The  condition  frequently  encountered  in  the 
Lepidoptera  is  that  of  a  distinct  separation  of  the  tergum  and  sternum  of  segment  IX. 

The  valva  is  approximately  triangular,  and  unmodified.  This  condition  is  found  in  many  species  of 
macrolepidopterans  including  many  butterflies. 

There  are  no  particularly  striking  modifications  of  the  main  components  of  the  genitalia.  The  greatest 


276 


M.  J.  SCOBLE 


Figs  72-76  Male  genital  capsules  of  Hedylidae.  72-74,  lateral  aspects  of  (72)  Macrosoma  tipulata 
Hiibner,  (73)  M.  lamellifera  (Prout),  (74)  M.  napiaria  (Guenee);  75,  76,  ventral  aspects  of  (75)  M. 
napiaria  (Guenee);  76;  M.  nigrimacula  (Warren)  (a)  valvae  and  juxta  removed,  (b)  valvae  and  juxta. 


modifications  are  the  shapes  of  the  uncus  in  heliconiaria  (Fig.  83),  the  valva  (Fig.  79)  and  juxta  (Fig.  84)  of 
hyacinthina,  and  the  simplification  of  the  genitalia  of  tipulata  (Fig.  72). 

Female  postabdomen  and  genitalia  (Figs  90-99).  Tergum  VII  is  longer  than  tergum  VI  and  forms  a  weak 
hood  over  the  genitalia,  when  they  are  not  extended.  Tergum  VIII  is  well-sclerotized  and  narrow  (Fig.  90), 
and  gives  rise  to  a  pair  of  thin,  anterior  apophyses.  In  most  species  examined  these  are  sclerotized,  but  in 
some  they  are  membranous  (staining  in  Chlorazol  black  E),  and  short  (Figs  93,  96).  Presumably,  in  those 
species  with  reduced  apophyses  the  muscles  are  functionless  during  oviposition.  In  hyacinthina  and 


NEW  CONCEPT  OF  BUTTERFLIES 


277 


78 


Figs  77-84  Male  genitalia  of  Hedylidae.  77-82,  valvae  of  (77)  Macrosoma  heliconiaria  (Guenee),  (78) 
M.  semiermis  (Prout),  (79)  M.  hyacinthina  (Warren),  (80)  M.  lamellifera  (Prout)  (comb,  n.),  (81)  M. 
nigrimacula  (Warren),  (82)  M.  napiaria  (Guenee);  83,  uncus  of  M.  heliconiaria  (Guenee),  ventral  view; 
84,  juxta  of  M.  hyacinthina  (Warren),  valvae  at  different  plane  of  focus. 


278 


M.  J.  SCOBLE 


Figs  85-89     Male  genitalia,  aedeagus,  of  Hedylidae.  85,  Macrosoma  heliconiaria  (Guenee);  86,  M. 
hyacinthina  (Warren);  89,  M.  napiaria  (Guenee). 


hedylaria  the  reduced  anterior  apophyses  are  usually  bent  inwards  (medially)  at  their  ends  at  about  right 
angles.  Reduced  anterior  apophyses  are  found  in  many  butterflies. 

Terga  IX  and  X  form  a  sclerotized  band  more  narrow  than  tergum  VIII.  The  posterior  apophyses,  which 
are  derived  from  this,  are  sclerotized  rods. 

Ventrally,  sternum  VII  is  folded  before  the  ostium  bursae  thus  forming  a  lip  to  this  aperture.  The  lip  is 
nearly  always  fringed  with  strong  setae  (Fig.  90).  The  sclerotizations  around  the  ostium  bursae  include 
anterior,  posterior,  and  lateral  components.  The  antevaginal  and  postvaginal  sclerites  are  small,  while  the 
lateral  sclerites  (derived  from  sternum  VIII)  are  large  and  extend  laterally  to  unite  with  the  anterior 
apophyses  in  those  species  where  they  are  sclerotized.  The  extent  of  these  ostial  sclerotizations  varies 
between  species.  In  heliconiaria  the  lamella  postvaginalis  is  absent  as  are  the  setae  on  the  fold  of  sternum 
VII.  In  tipulata  the  sclerotizations  are  virtually  absent  (Fig.  97). 


NEW  CONCEPT  OF  BUTTERFLIES 


279 


91 


Figs  90-91     Female  postabdomen  and  genitalia  of  Hedylidae.  90,  Macrosoma  hedylaria  (Warren), 
abdominal  sternum  VII,  ventral  aspect,  antrum  (an);  91,  M.  lucivittata  (Walker),  lateral  aspect. 


Figs  92-95  Female  genitalia  of  Hedylidae.  92-94,  ventral  aspect  of  (92)  Macrosoma  rubedinaria 
(Walker),  (93)  M.  hyacinthina  (Warren),  anterior  apophysis  arrowed,  (94)  M.  rubedinaria  (Walker)  to 
show  well-sclerotized  anterior  apophyses;  95,  M.  lucivittata  (Walker),  lateral  aspect. 


280 


M.  J.  SCOBLE 


98 


Figs  96-99  Female  genitalia  of  Hedylidae.  96,  97,  lateral  aspect  of  (96)  Macrosoma  hedylaria  (Warren), 
anterior  apophysis  arrowed,  (97)  M.  tipulata  Hiibner;  98, 99,  signum  of  (98)  M.  lucivittata  (Walker),  (99) 
M.  rubedinaria  (Walker). 


The  ostium  bursae  leads  into  the  ductus  bursae,  the  first  part  of  which  is  funnel-shaped  with  sclerotized 
walls,  the  antrum  (Fig.  94).  Beyond  this  the  ductus  bursae  is  narrow,  membranous,  and  usually  long.  It 
expands  suddenly  into  a  globose  or  slightly  elongate-globose  membranous  corpus  bursae  (Fig.  91).  The 
corpus  bursae  may  bear  a  single  signum  of  a  characteristic  shape  (Figs  98, 99) ,  or  the  signum  may  be  absent. 

The  weakly  telescoped  ovipositor  ends  in  a  pair  of  large,  soft,  setose,  ear-like  lobes  (Figs  92,  93). 

Juvenile  stages 

The  shape  of  the  egg  varies  within  the  Lepidoptera,  but  it  is  essentially  flat  or  upright  (Chapman,  1896). 
Many  butterflies  have  upright  eggs.  In  the  Hesperiidae,  however,  they  are  flat.  The  oval  egg  dissected  from 
the  macerated  abdomen  of  a  female  hedylid  appears  to  be  of  the  flat  variety,  but  its  shape  did  not  resemble 
that  of  the  hesperiid  egg.  Since  both  flat  and  upright  eggs  are  found  within  single  families,  or  even  single 
genera,  (Hinton,  1981),  the  gross  shape  of  the  egg  is  of  limited  phylogenetic  significance. 

The  larva  (Fig.  19)  exhibits  features  of  various  butterfly  families  rather  than  any  one  of  them.  Its 
horn-like  processes  (Figs  19, 100, 101)  strongly  resemble  those  of  many  Nymphalidae.  The  secondary  setae 
on  the  body  are  like  those  of  some  pierids.  The  crochets  on  the  ventral  prolegs  of  the  final  instar ,  which  are 
arranged  in  a  penellipse  or  as  transverse  bands,  are  neither  like  those  of  hesperiids  (in  which  they  form  a 
circle)  nor  like  papilionoids  (where,  in  post  first  instars,  they  are  usually  arranged  in  a  mesoseries) .  An  anal 
comb  (found  in  Hedylidae)  is  usually  present  in  Hesperiidae  and  at  least  some  Pieridae  among  the 
macrolepidopterans. 

The  pupa  (Figs  20, 21)  resembles  that  found  in  Papilionidae  (see  Igarashi,  1984)  or  Pieridae,  amongst  the 
butterflies,  in  that  it  is  attached  to  the  substrate  by  both  a  girdle  and  a  cremaster,  and  that  there  is  no 
cocoon.  A  girdle  is  present  in  some  sterrhine  geometrids,  e.g.  Anisodes  Guenee  (see  Common,  1986)  and 
Cyclophora  Hiibner.  However,  these  genera  lack  the  other  butterfly  characters  discussed  in  the  present 


NEW  CONCEPT  OF  BUTTERFLIES 


281 


100 


Figs  100,  101     Half  of  head  capsule  of  larval  exuviae  of  Hedylidae.  100,  Macrosoma  heliconiaria 
(Guenee);  101,  M.  nigrimacula  (Warren). 


work.  Furthermore,  whereas  the  girdles  of  butterflies  (including  hedylids)  are  thoracic,  those  of  sterrhine 
geometrids  are  spun  around  the  abdomen  (Holloway,  pers.  comm.  -  information  derived  from  unpub- 
lished manuscripts  housed  in  the  British  Museum  (Natural  History);  and  see  Fig.  3  of  Common,  1986  for  an 
Anisodes  girdle).  The  presence  of  a  thoracic  girdle  is  probably  a  ground  plan  character  of  the  Rhopalocera, 
although  weak  cocoons  are  usually  present  in  hesperiids.  Loss  of  the  cocoon  is  a  typically  papilionoid 
character,  although  it  is  present  in  Parnassius  Latreille  (Papilionidae)  and  weak  and  web-like  in  some 
satyrines.  The  girdle  is  present  in  Pieridae  and  Papilionidae,  but  is  lost  in  Nymphalidae  and  many 
Lycaenidae. 

A  termporal  cleavage  line  is  said  by  Scott  (1985)  to  be  present  in  Hesperiidae,  but  absent  from 
Papilionoidea.  Mosher  (1916)  referred  to  the  line  as  the  epicranial  suture.  The  apparent  absence  of  the  line 
in  the  Hedylidae  is  therefore  like  that  of  true  butterflies  rather  than  that  of  skippers.  A  temporal  cleavage 
line  is  found  in  many  moths.  Mosher  notes  its  presence  in  Lycaenidae,  although  Scott  (1985:  table  1) 
records  it  as  absent  from  that  family. 

Discussion 

The  Hedylidae  exhibit  a  high  degree  of  structural  uniformity.  Several  characters  seem  to  be 
unique  to  the  family,  which  provides  evidence  for  its  suspected  monophyly.  The  most  striking 
are  the  small  chambers  at  the  base  of  the  fore  wing,  the  protuberances  on  the  underside  of  the 
hind  wing,  the  sinuous  course  of  veins  Rsi  and  Rs2  of  the  radial  sector  in  the  fore  wing,  and  the 
long  fore  tarsi  of  the  male  composed  of  only  two  tarsomeres. 

Two  important  questions  about  the  phylogeny  of  the  Hedylidae  need  to  be  discussed.  First, 
are  hedylids  members  of  a  taxon  composed  of  Hesperioidea  plus  Papilionoidea  plus  themselves, 
i.e.,  are  they  butterflies  in  the  broadest  sense?  Second,  if  so,  what  are  the  relationships  of 
hedylids,  hesperiids,  and  papilionoids  to  each  other:  do  hedylids  represent  the  sister  group  of  the 
taxon  Hesperioidea  plus  Papilionoidea  (i.e.  the  Rhopalocera),  or  are  they  more  closely  related 
to  the  Hesperioidea  or  to  the  Papilionoidea  than  are  either  of  these  two  related  to  each  other?  I 
shall  argue  that  the  Hedylidae  are  indeed  'butterflies'  in  the  broadest  sense.  That  they  may 
represent  the  sister  group  of  the  Papilionoidea  remains  an  intriguing  possibility.  This  possibility 
is  not  a  conclusion  of  this  work:  it  would  be  premature  and  requires  further  comparison  of  the 
three  taxa  involved. 


282  M.  J.  SCOBLE 

A  separate,  but  related,  problem  is  how  to  treat  the  nomenclature.  Papilionoidea  and 
Hesperioidea  are  widely  used  names,  although  Hesperioidea,  consisting  as  it  does  of  a  single 
family,  is  redundant  (Farris,  1976;  Wiley,  1979),  assuming  that  the  Megathyminae  are  not 
viewed  as  an  independent  family.  How  should  the  Hedylidae  be  treated? 

Although  the  butterflies  are  one  of  the  best  studied  groups  of  organisms,  the  published 
evidence  for  their  monophyly  is  certainly  not  overwhelming.  Ehrlich  (19586)  regarded  hesper- 
iids  as  papilionoids  that  retain  a  great  many  primitive  characters.  Both  Ehrlich  and  Kristensen 
(1976)  retained  the  two  superfamilies  and  did  not  synonymize  them,  Ehrlich  because  he 
considered  the  phenetic  distance  between  them  sufficiently  great,  and  Kristensen  because  he 
considered  the  sister-group  relationship  a  reasonable  working  hypothesis  but  one  that  required 
confirmation.  Although  the  consensus  is  that  hesperioids  and  papilionoids  do  form  a 
monophyletic  group,  the  characters  on  which  this  argument  is  based  are  not  entirely  convincing. 

Kristensen  (1976)  records  six  possible  specialized  characters  (apomorphies)  shared  by  the  two 
taxa,  two  of  which  he  considers  doubtful.  However,  none  of  them  have  been  examined  in  a  great 
number  of  species  of  butterflies  and  moths.  Until  they  are  examined  they  must  remain  somewhat 
in  doubt.  A  particular  problem  with  these  characters  is  that  they  are  not  observable  on  dried 
specimens.  Some  of  the  characters  can  only  be  seen  on  alcohol-preserved,  or  fixed  material  (e.g. 
the  twist  in  the  oblique  lateral  dorsal  muscle  of  the  mesothorax,  and  the  structure  of  the  aorta  in 
the  mesothorax).  Those  that  can  be  observed  on  dried  specimens  require  maceration  of  the  head 
or  thorax,  and  have  been  examined,  inevitably,  in  a  limited  number  of  Lepidoptera.  A  detailed 
study  of  the  relationships  of  the  Hesperioidea  and  the  Papilionoidea  is  being  undertaken  by  Dr 
R.  de  Jong,  who  has  found  several  specialized  characters  considered  to  support  the  sister-group 
relationship  of  the  two  taxa.  If  hedylids  are  the  sister  group  of  papilionoids  then  several  of  de 
Jong's  characters  will  have  to  be  treated  as  homoplasious.  It  would  be  premature  to  treat  the 
Hedylidae  as  the  sister  group  of  the  Papilionoidea  while  de  Jong's  detailed  work  is  in  progress. 
For  this  reason  it  has  not  been  possible  to  make  unequivocal  statements  about  precisely  which 
taxa  some  of  the  characters  'define'. 

Recently  Scott  (1985)  has  discussed  the  phylogeny  of  the  butterflies.  From  his  table  1,  the 
characters  that  appear  to  support  the  monophyly  of  the  papilionoids  and  hesperioids  are:  the 
presence  of  a  postspiracular  bar,  found  in  skippers  but  not  in  Papilionidae,  some  Nymphalidae, 
and  Lycaenidae,  and  the  presence  of  a  secondary  sternopleural  sulcus,  variable  in  size,  but 
present  in  all  butterflies. 

No  other  specialized  features  are  tabulated  by  Scott  as  unique  to  the  adults  of  all  butterflies, 
with  the  possible  exception  of  the  absence  of  spurs  on  the  tibia  of  the  mid  leg.  However, 
although  the  spurs  are  absent  from  skippers,  they  are  lost  only  in  Papilionidae  within  the  true 
butterflies  so  the  condition  is  likely  to  be  a  parallel  development. 

Of  the  attributes  of  the  Hedylidae  described  and  discussed  above,  13  are  of  particular 
significance  in  the  discussion  of  the  question  of  the  relationship  of  the  Hedylidae  to  the  skippers 
and  true  butterflies.  Of  these  several  resemble  the  condition  in  true  butterflies  rather  than 
skippers. 

(1).  Apophyses  of  metathoracic  furca  sagittate.  The  sagittate  condition  appears  to  be  a 
specialized  character  of  the  three  taxa.  It  certainly  deserves  examination  in  many  more  species 
to  establish  that  it  is  definitely  absent  from  moths  and  therefore  diagnostic  of  butterflies. 

(2).  Pupa  girdled.  Ehrlich  (19586)  and  Scott  (1985)  regard  a  girdled  pupa  in  butterflies  as 
primitive  and  its  absence  as  specialized.  Scott  considers  this  condition  to  have  been  present  in 
the  ancestor  of  hesperioids  and  papilionoids.  There  is  no  general  survey  of  the  occurrence  of 
girdled  pupae  in  the  Lepidoptera,  so  to  suggest  that  the  condition  is  diagnostic  of  hedylids, 
hesperioids,  and  papilionoids  needs  to  be  treated  with  caution.  This  attribute  derives  its 
taxonomic  strength  in  conjunction  with  others. 

(3).  Second  median  plate  of  fore  wing  lies  partly  under  the  base  of  vein  L4.  Although  this 
character  requires  examination  in  a  wide  range  of  Lepidoptera,  it  appears  to  support  the  view  of 
the  monophyly  of  the  Hesperioidea  plus  the  Papilionoidea  (de  Jong,  pers.  comm.)  plus  the 
Hedylidae. 

(4).  Presence  of  an  anal  comb  in  the  larva.  This  structure,  which  is  present  in  Hesperioidea 


NEW  CONCEPT  OF  BUTTERFLIES  283 

and  Hedylidae,  seems  to  be  confined  to  the  Pieridae  in  the  Papilionoidea.  It  may  reasonably  be 
assumed  to  be  a  ground  plan  character  of  the  Papilionoidea.  The  presence  of  an  anal  comb  in  the 
Tortricidae  is  probably  an  independent  development.  However,  if  the  Macrolepidoptera  are  not 
monophyletic  then  the  closest  relatives  of  the  Rhopalocera  may  be  found  among  the  microlepi- 
dopterans.  The  tortricoid/cossoid  assemblage  might  be  a  serious  contender. 

(5).  Presence  of  a  postspiracular  bar  on  the  first  abdominal  segment.  Postspiracular  scler- 
otizations  are  present  on  the  first  abdominal  segment  of  non-ditrysian  moths  (see  for  example 
Kyrki,  1983;  Kristensen,  1984a  -  'lateral  lobe  of  tergum  F  lo  in  his  figure  2).  That  the  presence  of 
a  secondary  sclerotization  (postspiracular  bar)  is  a  character  of  the  ground  plan  of  hedylids, 
hesperioids,  and  papilionoids  is  a  reasonable  supposition.  The  presence  of  a  postspiracular  bar 
in  certain  Pyraloidea  (those  without  tympanal  organs),  certain  Geometroidea  (Drepanidae, 
Thyatiridae,  and  most  Uraniidae),  and  Noctuoidea  (many  families)  (Brock,  1971)  must  cast 
some  doubt  on  the  value  of  the  structure  as  an  indicator  of  the  monophyly  of  hedylids, 
hesperioids,  and  papilionoids.  The  postspiracular  bar  is  recorded  as  absent  from  macrolepidop- 
terans  by  Scott  (1985:  table  1),  although  he  made  no  comment  on  the  postspiracular  sclerotiza- 
tions  in  the  macrolepidopteran  families  discussed  by  Brock  (1971). 

(6).  Reduction  of  anterior  apophyses  in  female  genitalia.  The  weak,  reduced  anterior 
apophyses  in  some  Hedylidae  is  a  condition  found  also  in  many  butterflies  (de  Jong,  pers. 
comm.).  Whether  this  is  a  ground  plan  character  of  hedylids,  hesperioids,  and  papilionoids,  or 
whether  it  is  one  that  is  subject  to  extensive  parallelism  is  unknown. 

(7).  Abdomen  curved,  particularly  in  males.  As  noted  above,  the  shape  of  the  abdomen 
resembles  that  of  the  true  butterflies  rather  than  that  found  in  skippers.  This  character  is  only 
doubtfully  a  unique  derivation  of  Hedylidae  +  Papilionoidea.  A  detailed  study,  preferably 
measuring  the  degree  of  curvature,  is  needed  to  ascertain  whether  the  condition  in  Hedylidae 
and  Papilionoidea  is  greater  than  that  found  in  Hesperiidae. 

(8).  Abdominal  tergum  I  strongly  'pouched'. 

(9).  Precoxal  (paracoxal)  sulcus  joins  'marginopleural'  sulcus.  This  character  appears  to 
be  a  specialization  of  the  Hedylidae  and  the  Papilionoidea.  The  main  objections  to  this  view 
are  that  the  sulci  are  difficult  to  observe,  and  that  they  have  been  examined  in  relatively  few 
species. 

(10).  Pupal  cocoon  lost.  Unlike  the  condition  in  hesperioids,  the  cocoon  of  hedylids  and  most 
papilionoids  is  lost.  Loss  of  a  cocoon  occurs  elsewhere  in  the  Lepidoptera,  so  this  character  is  of 
doubtful  value  and  derives  its  strength  in  consideration  with  others. 

(11).  Loss  of  temporal  cleavage  line  in  pupa.  The  loss  of  this  cleavage  line  in  both  hedylids 
and  papilionoids  is  further,  although  rather  weak,  evidence  for  their  close  relationship. 

(12).  Crochets  of  ventral  prolegs  of  larva  not  forming  a  complete  circle.  This  character  is 
based  on  an  examination  of  only  two  species.  The  penellipse  ofnigrimacula,  and  the  transverse 
bands  of  heliconiaria  are  neither  the  typical  circle  of  hesperiids  nor  the  usual  mesoseries  of 
papilionoids.  It  is  possible,  considering  other  characters  of  the  family,  that  the  hedylid  condition 
is  part  of  a  transformation  series  circle  — >  penellipse  — >  mesoseries,  but  this  remains  a  tentative 
suggestion. 

(13).  Loss  of  pretarsus  in  the  fore  leg  of  males.  Reduction  or  loss  of  the  pretarsus  is 
widespread  in  butterflies.  A  reduction  of  the  pulvilli  and  arolium  is  the  condition  found  in  all  legs 
of  the  most  primitive  papilionoids.  The  initial  reduction,  confined  as  it  is  to  the  fore  legs  of  the 
male,  appears  to  be  the  first  stage  of  a  reduction  fully  established  in  the  papilionoids.  Therefore 
either  the  character  is  a  shared  specialization  of  the  Hedylidae  plus  Papilionoidea  or  it  has 
developed  in  parallel  in  the  two  groups.  The  possibility  remains  that  the  pretarsus  is  redeveloped 
(a  reversal)  in  hesperiids. 

Nomenclature 

The  Papilionoidea  are  generally  regarded  as  a  separate  superfamily  from  the  Hesperioidea,  although 
Brock  (1971)  combined  the  two  in  Papilionoidea.  In  the  present  study  I  follow  convention  and  retain  both 
taxa  as  superfamilies  since  the  relationships  between  the  Hesperioidea  and  the  Papilionoidea  is  being 
studied  in  detail  by  R.  de  Jong.  Also,  Kristensen  (1976)  noted  some  uncertainties  about  the  sister  group 


284  M.  J.  SCOBLE 

relationship  of  the  two  groups,  and  recently  Stallingseffl/.  (1985,  quoting  C.  L.  Remington)  made  a  similar 
point. 

It  would  be  premature  to  formally  asign  the  Hedylidae  to  the  Papilionoidea  at  this  stage,  so  to  retain 
equivalence  of  rank  they  are  treated  as  a  superfamily  (Hedyloidea  stat.  n.).  Consequently,  the 
Rhopalocera  now  include  three  superfamilies  Hesperioidea,  Hedyloidea,  and  Papilionoidea,  all  of 
interchangeable  position  (sedis  mutabilis,  Wiley,  1979). 

Epilogue:  a  suggested  phytogeny 

Although  this  work  concluded  with  the  conservative  suggestion  that  hedylids  are  rhopalocerans 
of  which  the  precise  relationships  remain  uncertain,  the  following  dendrogram  (Fig.  102)  is  a 
fully  resolved  three  taxon  statement  presented  to  stimulate  critical  assessment. 


7-13 


1-6 


Fig.  102  Dendrogram  to  suggest  possible  phylogenetic  relationships  of  Hesperioidea,  Hedyloidea  and 
Papilionoidea.  Numbers  represent  the  characters  in  the  Discussion.  Relative  strengths  and  weaknesses 
of  the  characters  are  not  indicated,  but  are  considered  in  the  text. 

Acknowledgements 

I  am  indebted  to  my  colleagues  for  their  interest  in  this  work,  and  for  their  critical  comments  and 
suggestions  throughout  its  course.  In  particular  I  thank  P.  R.  Ackery,  D.  J.  Carter,  J.  D.  Holloway,  I.  J. 
Kitching,  K.  Sattler,  J.  A.  Schoorl,  R.  I.  Vane-Wright,  A.  Watson  and  P.  E.  S.  Whalley.  I  am  grateful  to 
M.  J.  Matthews,  who  helped  to  record  distribution  data.  Useful  information  was  also  received  from  V.  O. 
Becker.  R.  O.  Kendall  supplied  Juvenile  stages  of  Macrosoma  heliconiaria,  which  added  an  extra 
dimension  to  this  study.  I  am  extremely  grateful  to  him.  R.  de  Jong  and  N.  P.  Kristensen  provided  very 
valuable  comments  on  the  manuscript.  I  thank  both,  and  further  express  my  gratitude  to  R.  de  Jong  who 
made  available  unpublished  sketches  and  other  information  from  his  work  on  the  phylogenetic  rela- 
tionships within  the  butterflies.  I  thank  T.  Fox  and  D.  Lees  for  generously  supplying  collection  details, 
photographs  of  a  larva  and  adult,  and  pupal  exuviae  of  M.  cascaria.  I  am  grateful  to  members  of  the 
Photographic  Unit  of  the  British  Museum  (Natural  History)  for  many  of  the  illustrations:  Ms  S.  H.  Barnes, 
scanning  electron  micrographs;  P.  V.  York,  photo  micrographs;  H.  Taylor,  photo  macrographs. 

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81-93. 


British  Museum  (Natural  History) 

Milkweed  butterflies:  their  cladistics  and  biology 

P.  R.  Ackery  &  R.  I.  Vane- Wright 

The  Danainae,  a  subfamily  of  the  Nymphalidae,  contains  only  some  150  species,  yet  aspects  of 
their  biology  have  stimulated  far  more  attention  than  can  be  justified  by  species  numbers 
alone.  In  recent  years,  an  expansive  literature  has  grown,  considering  aspects  of  their 
courtship  and  pre-courtship  behaviour,  migration,  larval  hostplan  associations,  mimicry  and 
genetics.  The  popularity  of  danaines  among  biologists  can  certainly  be  attributed  to  this 
combination,  within  one  small  group,  of  so  many  of  the  factors  that  make  butterflies  such  an 
interesting  group  to  study.  The  obvious  need  to  place  this  wealth  of  biological  data  within  an 
acceptable  systematic  framework  provided  the  impetus  for  this  volume. 

Started  eight  years  ago  within  the  conventions  of  evolution  by  natural  selection  and 
Hennig's  phylogenetic  systematics,  the  book  is  now  largely  about  natural  history  (what  the 
animals  have  and  do,  where  they  live  and  how  they  develop)  and  natural  groups  -  as  revealed 
by  a  form  of  analysis  approaching  that  practised  by  the  new  school  of  'transformed  cladistics' . 
The  authors  have  prepared  a  handbook  that  will  appeal  to  a  wide  range  of  biologists,  from 
museum  taxonomists  to  field  ecologists. 

425  pp,  12  pp  colour,  73  b/vv  plates,  line  and  graphic  illustrations,  maps,  extensive  bibliography. 
ISBN  0  565  00893  5. 1984.  Price  £50. 


Titles  to  be  published  in  Volume  53 


A  review  of  the  Miletini  (Lepidoptera:  Lycaenidae) 

By  J.  N.  Eliot 

Australian  ichneumonids  of  the  tribes  Labenini  and  Poecilocryptini 

By  I.  D.  Gauld  &  G.  A.  Hollo  way 

The  tribe  Pseudophloeini  (Hemiptera:  Coreidae)  in  the  Old  World  tropics  with  a  discussion  on 
the  distribution  of  the  Pseudophloeinae 

By  W.  R.  Dolling 

The  songs  of  the  western  European  grasshoppers  of  the  genus  Omocestus  in  relation  to  their 
taxonomy  (Orthoptera:  Acrididae) 

ByD.  R.  Ragge 

The  structure  and  affinities  of  the  Hedyloidea:  a  new  concept  of  the  butterflies 

ByM.  J.  Scoble 


Photoset  by  Rowland  Phototypesetting  Ltd,  Bury  St  Edmunds,  Suffolk 
Printed  in  Great  Britain  by  Henry  Ling  Ltd,  Dorchester 


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